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 Transition. Princeton, 1944. 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. no. 6, pp. 5-49. 1907. 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 the Compiler. Cambridge Univ. Press, 1940. *8tiUman Stillman, J. D. B., Seeking the Golden Fleece; a Record of Pioneer Life in California . . . , esp. pp. 285-326. San Francisco, 1877. *Stone Stone, Witmer, Philadelphia to the coast in early days, and the development, of western ornithology prior to 1850. Condor, 18:3-14, 1916. Sivainson Swainson, William, Taxidermy, with the Biography of Zoologists, and Notices of their Work. London, 1840. 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. Camp. Columbus, Ohio, 1953. Woodcock d Steam Woodcock, H. B. D., and William T. Stearn, Lilies of the World. London and New York, 1950. BIBLIOGRAPHY Adams, James Capen J. Grinnell, J. S. Dixon, and J. M. Linsdale, Fur-beaiing Mammals of California, 1:78-80, 1937. Agassiz, Alexander Emmanuel Rudolphe, 1835-1910 Carpenter, 2; DAB; Meisel, 1:156; A. G. Mayer in Pop. Sci. Mo., 77: 418-446 (portr.), 1910; G. R. Agassiz, Letters and Recollections of Alexander Agassiz, 161-162 et liossim. 1913; G. L. Goodale in Nat. Acad. Sci. Biog. Mem.. 7:291-305 (portr.), 1912. Agassiz, Jean Louis Rudolphe, 1807-1873 Carpenter, 2; DAB; Dall, imssim; Meisel, 1:157-158; Sherborn; D. S. Jordan in Science, n.s., 16:250, 1902, and Days of a Man, 1:107-117 (portr.), 1922; W. J. Youmans, Pioneers of Science in America, 475-491 (portr.), 1896. Allen, Charles Andrew, 1841-('?) Bade, 2:71, but perhaps not this person (?); Palmer. 264; cf. Forest and Stream. 7:4, Aug. 10, 1876. Ame-s, Mary E. Pulsifer, 1845-1902 Brewer, 558; Boston Evening Transcript for Mar. 21, 1902. EWAN: SAN FRANCISCO AS A MECCA FOR NINETEENTH CENTURY NATURALISTS 43 Anderson, Charles Lewis, 1827-1910 Brewer, 558; Jepson, 1:214-216 (portr.), 1929; Wagner-three, 219a. Andersson, Nils Johan, 1821-1880. Breioer, 557; Rydherg, 25; VrMn Fl. Bras., 1:1; Yan Steenis 16. Andrews, Timothy Langdon, 1819-1908 Brewer, 557; Howell, 30; L. M. Pammel in Cedar Rapids [Iowa] Daily Republican for Mar. 19, 1908 (portr.). AuDXJBON, John Woodiiouse, 1812-1862 Dall, 155 et passim; Geiser-two. 270; Meisel, 1:161; Wagner-three, 176, 208; H. Harris in Condor, 43:34, 1941; Maria R. Audubon, Audubon and His Journals, ed. by E. Coues, passivi, 1898. Austin, Rebecca Merritt Smith Leonard, 1832-1919 Brewer, 558; Jepson, 2:130-133 (portr.), 157; M. A. H[ail] [daughter of Mrs. Austin] in Plumas [County] National Bulletin, vol. 53 no. 29, for Mar. 27, 1919. copy examined in Frank Morton Jones Library, Wilmington, Del.; H. S. Reed in Oakland [Calif.] Tribune for Dec. 28, 1941; F. M. Jones in Delaware Notes, ser. 23:24-35, 1950. Ayres, William Orville, 1817-1891 Dean, 1:45; Dall, 154; Farquhar's Yosemite, 11, 26, etc., for Thomas A. Ayres, whose relationship to Wm. 0. Ayres is undetermined; cf. A. Kellogg in Proc. Calif. Acad. Sci. for Feb. 3, 1873, for proposed genus Ayresia. Baker, Milo Samuel, 1868- Eioan, 155; cf. Madrono, 4:283, 1938. Barclay, George Brewer, 555; Britten. 19; cf. Palmer. 268 s.v. Belcher; Van Steenis. 36. Barkelew, Frederick E. Cf. E. W. Nelson in Mem. Nat. Acad. Sci., 16:144, 1921; I. M. Johnston in Proc. Calif. Acad. Sci., ser. 4, 20:13. 1931; P. A. Munz in Leafl. West. Bot.. 7:73, 1953. Barlow, Chester, 1874-1902 Palmer, 267; W H. Osgood in Auk, 20:92-93, 1903; H. R. Taylor in Condor, 5:3-7 (portr.), 1903. Beardsley, a. F. Brewer, 557; Farquhar's Brewer, 218-219; R. C. Miller in Calif. Hist. Soc. Quart.. 21:366, 1942, as "A. F. Beardslee," quoted from Academy's Proc. Beck, Rollo Howard, 1870- Palvier, 267-268; R. C. Murphy, Oceanic Birds of South America, 2:2.j (portr., pi. 1), 1936. Beechey, Frederick William, 1796-1856 Britten; DNB ; Stillman, 319-325. Behr, Hans Hermann, 1818-1904 Brewer, 556; Britten, 339; Carpenter, 7; Essig, 553-556 (portr.) ; cf. Geiser-tico. 271; Meisel, 1:164; anon, in Entom. News, 15:142-144, 1904 (from San Francisco Chronicle); A. Eastwood in Science, n.s., 19:636, 1904; autobiog. notes in Erythea, 4:168-173, 1896; A. E. Zucker, Forty-eighters, 277, 1950. 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. C(hapman) in Auk, 7:98-99, 1890; J. F. McDermott, Up the Missouri with Audubon, passim, 1951. BiDWELL, Annie E. Kennedy (Mrs. John Bidwell) Bade, 2:72 et passim; R. D. Hunt, John Bidwell, Prince of California Pioneers, Cax- ton Press, Caldwell, Idaho, 1942; D. S. Jordan, Days of a Man, 1:471, 1922. 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. REFERENCES Andrewes, C. H. 1952. Classification and nomenclature of viruses. Ann. Rev. Microbiol., 6:119-138. Andrewes, F. W. 1930. The nomenclature and classification of micro-organisms. In A System of Bac- teriology, 1:292-310. London: Med. Res. Council. VAN NIEL: SYSTEMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 1 1 1 Babcock, E. B. and G. L. Stebbins, Jk. 1938. The American Species of Crepis. Carnegie Inst. Washington Publ. 504. 199 pp. Washington. Badian, J. 1933. Eine cytologische Uutersuchuug iiber das Chromatin und den Entwicklungs- zyclus der Bakterien. Arch. Mikrobiol., 4:409-418. Beijebinck, M. W. 1921. Verzamelde Geschriften. Vols. IV. The Hague: M. Nyhoff. Benecke, W. 1912. Ban und Leben der Bakterien. xii + 650 pp. Leipzig: Teubner. BissETT, K. A., J. Greene, and E. 0. Morris 1951. The nuclear reduction process in bacteria. Exp. Cell. Res., 2:388-392. BoiviN, A. 1948. Les acides nucleiques dans la constitution cytologique et dans la vie de la cellule bacterienne. Compt. Rend. Soc. Biol., 142:1258-1275. Breed, R. S., E. G. D. Murray, and A. P. Hitchcns, eds. 1948. Bergey's Manual of Determinative Bacteriology. 6th ed. xvi + 1529 pp. Bal- timore: Williams and Wilkins Co. Buchanan, R. E. 1916-1918. Studies in the nomenclature and classification of the bacteria. I-X. Journ. Bact., 1(1916) :591-596; 2(1917) :155-162, 347-350, 603-617; 3(1918): 27-61, 175-181, 301-306, 403-406, 461-474, 541-545. 1925. General Systematic Bacteriology. 597 pp. Baltimore: Williams and Wilkins Co. BUDEE, J. 1919. Zur Biologie des Bakteriopurpurins und der Purpurbakterien. Jahrb. Wiss. Eot., 58:525-628. BUTSCHXI, O. 1889. Protozoa. In H. G. Bronn's Klassen und Ordnungen des Thier-Reichs. Vols. I-III. Leipzig and Heidelberg: C. F. Winter. Calvin, M., and V. Lynch 1952. Grana-like structures of Synec?iococcus cedorum. Nature, 16g: 455-456. Camp, W. H. 1951. Biosystematy. Brittonia, 7:113-127. Camp, W. H., and C. L. Gilly 1943. The structure and origin of species. Brittonia, 4:323-385. COHN, F. 1854. Untersuchungen iiber die Entwicklungsgeschichte der mikroskopischen Algen und Pilze. Nov. Act. Leo.-Carol., 24:103-256. 1872. Untersuchungen iiber Bacterien. Beitr. Biol. Pflanz., Heft 2, pp. 127-224. 1875. Untersuchungen uber Bacterien, II. Beitr. Biol. Pflanz., Heft 3, pp. 141 208. Copeland, H. F. 1938. The kingdoms of organisms. Quart. Rev. Biol., 13:383-420. 1947. Progress report on basic classification. Amer. Nat. 81:340-350. DeLamater, E. D. 1952. Preliminary observations on the occurrence of a typical mitotic process in micrococci. Bull. Torrey Club, 79:1-5. 112 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES DeLamater, E. D. (Cont.) 1952. Evidence for the occurrence of true mitosis in bacteria and certain applica- tions. Journ. Invest. Dermatol., 18:225-230. Delapobte, B. 1939. Recherches cytologiques sur les Bacteries et les Cyanophycees. 160 pp. Paris: Andre Lesot. DOBZHANSKY, T. 1941. Genetics and the Origin of Species, xiii + 446 pp. New York: Columbia Univ. Press. Endeblein, G. 1925. Bakterien-Cyclogenie. 390 pp. Berlin and Leipzig: De Gruyter & Co. Geitlek, L. 1932. Cyanophyceae. In L. Rabenhorst, Kryptogamen-Flora von Deutschland, oster- reich und der Schweiz. 2d ed. Vol. XIV. 1196 pp. GiLMOUK, J. S. L. 1951. The development of taxonomic theory since 1851. Nature: 168:400-402. Haeckel, E. 1866. Generelle Morphologie der Organismen. Bd. I. Allgemeine Anatomie der Organismen. xxxii + 574 pp. Bd. II. Allgemeine Entwickelungsgeschichte der Organismen. clx + 462 pp. Berlin: Georg Reimer. Hayes, W. 1952. Recombination in Bad. coli K12. Unidirectional transfer of genetic material. Nature, 169:118-119. 1952. Genetic recombination in Bact. coli K12. Analysis of the stimulating effect of ultraviolet light. Nature, 169:1017-1018. Kent, W. Saville 1880-1882. A Manual of the Infusoria. Vols. I-III. London: David Bogue. Kluyveb, a. J. and C. B. van Niel 1936. Prospects for a natural system of classification of bacteria. Centr. Bakt., II. Abt, 94:369-403. Knaysi, G. 1951. The structure of the bacterial cell. In C. H. Werkman and P. W. Wilson, eds.; Bacterial Physiology, espec. pp. 31-37. New York: Acad. Press. Knaysi, G., and R. F. Bakee 1947. Demonstration with electron microscope of a nucleus in Bacillus mycoides grown in a nitrogen-free medium. Journ. Bact., 54:4-5. Lederberg, J. 1947. Gene recombination and linked segregations in Escherichia coli. Genetics, 32:505-525. Lederberg, J., E. M. Lederberg, N. D. Zinder, and E. R. Lively 1951. Recombination analysis of bacterial heredity. Cold Spring Harbor Symp., 16:413-443. Lederberg, J., and E. L. Tatum 1946. Novel genotypes in mixed cultures of biochemical mutants of bacteria. Cold Spring Harbor Symp., 11:113-114. Lehmann, K. B., and R. 0. Neumann 1926-1927. Bakteriologie, insbesondere bakteriologische Diagnostik. 7th ed. Vols. I-II. Munich: J, F. Lehmann. VAN NIEL SYS7EMATICS OF THE BACTERIA AND BLUEGREEN ALGAE 1 ] 3 LlESKE, R. 1921. Morphologie und Biologie der Strahlenpilze (Actinomyceten). ix + 292 pp. Leipzig: Borntrager. MiGXJLA, W. 1897-1900. System der Bakterien. Vols. I-II. Jena: G. Fischer. 1907. Allgemeine Morphologie, Entwicklungsgeschichte, Anatomie und Systematik der Schizomyceten. In Lafar's Handbuch der technischen Mykologie, 1:29- 149. Jena: G. Fischer. MOLISCH, H. 1907. Die Purpurbakterien nach neuen Untersuchungen. vii + 95 pp. Jena: G. Fischer. Nageli, C. W. von 1857. Bericht Uber die Verhandlungen der botanischen Section der 33. Versamm- lung deutscher Naturforscher und Aertzte. Bot. Zeit., 15:760-761. NiEL, C. B. VAN 1931. On the morphology and physiology of the purple and green sulphur bacteria. Arch. Mikrobiol., 3:1-112. 1941. The bacterial photosyntheses and their importance for the general problem of photosynthesis. Advances in Enzymology, 1:263-328. 1944. The culture, general physiology, morphology, and classification of the non- sulfur purple and brown bacteria. Bact. Rev., 8:1-118. 1946. The classification and natural relationships of bacteria. Cold Spring Harbor Symp., 11:285-301. 1949a. The comparative biochemistry of photosynthesis. In J. Franck and W. Loomis, eds.. Photosynthesis of Plants, pp. 437-495. Ames, Iowa: Iowa State Coll. Press. 1949b. The "Delft School" and the rise of general microbiology. Bact. Rev., 13: 161-174. 1952. Bacterial Photosyntheses. In J. B. Sumner and K. Myrback, eds., "The En- zymes," 11:1074-1088. New York: Acad. Press. Oparin, a. I. 1938. The Origin of Life. (Transl. by S. Morgulis.) 270 pp. New York. Macmil- lan Co. Gela-Jensen, S. 1909. Die Hauptlinien des natiirlichen Bakteriensystems. Centr. Bakt., Abt. II. 22:305-346. Petbunkevitch, a. 1952. Maci"oevolution and the fossil record of Arachnida. Amer. Scient., 40:99-122. Piekakski, G. 1937. Cytologische Untersuchungen an Paratyphus- und Colibakterien. Arch. Mikro- biol., 8:428-439. POTTHOFF, H. 1922. Zur Entwicklungsgeschichte der Gattungen Chromatium und Spirillum. Centr. Bakt., Abt. II, 55:9-13. 1924. Zur Entwicklungsgeschichte der Bakteriengattungea Chromatium, SpiriUnm und Pseudomonas. Centr. Bakt., Abt. II, 61:249-255. Pbevot, a. R. 1933. fitudes de sj-st^matique bact^rienne. Ann. Sci. Nat. Bot, ser. 10, 15:23-260. 1940. Manuel de classification et de determination des bact^ries anaerobies. 223 pp. Paris: Masson et Cie. 114 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Pringsheim, E. G. 1949. The relationship between bacteria and Myxophyceae. Bact. Rev., 13:47-98. ROBINOW, C. F. 1944. Cytological observations on Bact. coli, Proteus vulgaris, and various aerobic spore-forming bacteria with special reference to the nuclear structures. Journ. Hyg., 43:413-423. 1945. Nuclear apparatus and cell structure of rod-shaped bacteria. In R. J. Dubos, The Bacterial Cell, "Addendum," pp. 355-377. Cambridge: Harvard Univ. Press. ScHACHMAN, H. K., A. B. PARDEE, and R. Y. Stanier 1952. Studies on the macromolecular organization of microbial cells. Arch. Biochem. and Biophys., 38:245-260. Skerman, V. B. D. 1949. A mechanical key for the generic identification of bacteria. Bact. Rev., 13:175-188. Stanier, R. Y. 1942. Agar-decomposing strains of the Actinomyces coelicolor species-group. Journ. Bact., 44:555-570. Stanier, R. Y., and C. B. van Niel 1941. The main outlines of bacterial classification. Journ. Bact., 42, 437-466. Stille, B. 1937. Zytologische Untersuchungen an Bakterien mit Hilfe der Feulgenschen Nu clealreaktion. Arch. Mikrobiol., 8:125-148. Tatum, E. L., and J. Lederberg 1947. Gene recombination in the bacterium, Escherichia coli. Journ. Bact., 53:673- 684. TlLASNE, R. 1947. Sur la mise en evidence du noyau des cellules bacteriennes. Compt. Rend. Soc. Biol., 141:411-413. 1949a. Sur la cytologie des bacteries vivantes etudiees grace au microscope a con- traste de phase. Compt. Rend. Soc. Biol., 143:1390-1392. 1949b. Donnees nouvelles sur la cytologie des bacteries apportees par la microscopie de contraste. Compt. Rend. Soc. Biol., 143:1392-1394. TuLASNE, R., and R. Minck 1947. Mise en evidence des noyaux des cocci par la ribonuclease. Compt. Rend Soc. Biol., 141:1255-1256. TuLASNE, R., and R. Vendrely 1947. Mise en evidence des noyaux bacteriens par la ribonuclease. Compt. Rend. Soc. Biol., 141:674-676. Welsch, M., and E. Nihoul 1948. A propos de la mise en evidence du noyau bacterien. Compt. Rend. Soc. Biol., 142:1449-1452. WlNOGRADSKY, S. 1949. Microbiologie du Sol. Oeuvres completes. 861 pp. Paris: Masson et Cie. 1952. Sur la classification des bacteries. Ann. Inst. Pasteur, 82:125-131. WiNSLOw, C. E.-A. 1914. The characterization and classification of bacterial types. Science, 39:77-91. AViN.'-LOW, C. E.-A., et al. 1917. Suggested outline of bacterial classification. Journ. Bact., 2:505-566. 1920. The families and genera of the bacteria. Journ. Bact. 5:191-229. 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 ply 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 of the Distoniatineae is essentially that of Senn (1900). He removed some of the forms which Klebs had placed in the Polymastigina to the Protomastigineae, abandoned the group Polymastigina, and elevated the Distomata to a group of major rank. (See Doflein, 1928, who retains the Polymastigina and credits it with seven families, p. 620.) The forms placed in this small group are characterized, among other fea- tures, by the double nature of the individuals — the body consisting of two halves and usually possessing two nuclei, two sets of flagella, and two oral fissures. (For a discussion on the occurrence of synzoospores in the algae in general, in- cluding the Chrysophyceae, and a comparison of them with representatives of the Distomatineae, reference should be made to two papers by Pascher: 1929, 1939.) All the representatives of the Distomatineae are placed in the family Disto- mataceae (Klebs) Blochmann orth. mut. Engler (1898, p. 7). CLASS BACILLARIOPHYCEAE Characterization: This class is comprised of uninucleate, diploid, unicellular and colonial, unattached (mostly free floating) or attached forms in which the inner part of the wall of the cell (known as the frustule) consists of pectin and the outer part of sili- ceous material. The wall is composed of two halves, one of which, the epitheca, is slightly larger and overlaps the other, the hypotheca. Each theca is in turn always composed of at least two pieces — the somewhat convex valve which is attached at its edges to the connecting band. It is the two connecting bands that overlap slightly, and together they constitute what is known as the girdle. In a number of forms the depth of the frustule is increased by the formation of one to many (depending upon the species) intercalary bands between the valves and their connecting bands. The valves are usually elaborately sculptured whereas the ornamentation in the con- necting bands is ordinarily much less conspicuous. The sculpturing is due to perforated thin areas (chambers) in the siliceous material (cf. Kolbe, 1948, pp. 4-12; Desikachary, PAPENFUSS: CLASSIFICATION OF THE ALGAE 151 1952) which appear as punctae or areolae. The striae of some Peiinales represent indi- vidual areolae or linear series of closely placed small areolae (punctae). With few excep- tions the markings on the two valves are similar. In many representatives of the order Pennales one or both the valves possess a com- plex system of slits and canals, the raphe system. Such forms are capable of independent gliding movement, apparently owing to cytoplasmic streaming. In colonial forms the cells are connected to one another in various ways by mucilage that is secreted through pores in the valves. Depending upon the genus, the cell contains one, two, or many yellow, olive-green, or brown chromatophores. Naked pyrenoidlike bodies are frequently present. The pig- ment complex consists of chlorophyll a. chlorophyll c, carotenes, and xanthophylls. Reserve food is stored as oil or leucosin. The usual method of reproduction is by vegetative cell division. The hypotheca of a dividing cell always becomes the epitheca of one of the two daughter cells. In the course of time there is thus in the vast majority of diatoms a considerable diminution in cell size in a population. Restoration of the maximal size characteristic of a species is brought about sooner or later by the production of rejuvenescent cells, called auxospores. In the majority of species investigated auxospores are formed as the result of a sexual process and two cells are usually involved. Meiosis precedes gametogenesis. One or at most two nonflagellated gametes are produced; or in certain Centrales four flagellated male gametes are formed (Stosch, 1951a). During the process of conjugation the gametes may escape from the parent frustules and fuse with those of the other cell. The zygote (auxospore) enlarges and ultimately produces a new frustule of maximal dimensions. Some species are autogamous and various instances of apogamy are on record. Various authors have reported the formation of small anteriorly or laterally biflagel- late cells (microspores) by members of the order Centrales. For a long time the function of these cells was unknown. Stosch (1951a, 1951b) and Geitler (1952) recently produced evidence that at least In some instances they are male gametes. Endogenous cysts with a wall consisting of two pieces, comparable to those of the Chrysophyceae and Xanthophyceae, have been observed in several members of the order Centrales. The Bacillariophyceae are divided into two orders, Centrales and Pennales, largely on the basis of the shape and ornamentation of the siliceous shell. In most Centrales, the valves are circular, angular, or irregular in outline and are radially or otherwise symmetrical with respect to a central point. In the Pennales the valves are isobilateral, medianly zygomorphic, or dorsiventral with at most only two planes of symmetry — one passing through the longitudinal axis, the other through the transverse axis, of the valve. In some members of this order the valves possess only one of these two planes of symmetry. Although a classification into two orders on this basis may seem to be artificial, it apparently is quite natural. In addition to the differences in the shape and ornamentation of the valves, the two orders differ in various other characters. Centric diatoms usually have many plastids, always lack a raphe and hence show no movement, produce cysts and may form motile male gametes. Pennate diatoms, on the other hand, usually have only one or two plastids, many possess a raphe and hence show movement, and do not form cysts or motile male gametes. History: The first person who described species of diatoms in a precise enough manner to afford their recognition by later workers was 0. F. Miiller. In 1773 he described a species of Gomphonema as Vorticella pyraria. Among the species which he described in later years, the most important is the one which he (0. F. Muller, 1783, p. 81, fn. c; 1786, p. 54) named Vihrio paxillifer. Gmelin (1788, p. 3903) established the genus BaciUaria for Miiller's Vibrio 2:)axillifer, and this name was later proposed by Nitzsch (1817), and following him used for a time especially by zoologists, for the entire group of so-called rod animalcules. The resemblance of many colonial diatoms to filamentous algae accounts for 152 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES the attention that these organisms received from various early botanists, who often described species under the generic name Conferva. Although as far as known, De Candolle did not especially investigate members of this group, he {in Lamarck and De Candolle, 1805, p. 48) was the first to regard the species previ- ously known by the name Conferva floccuJosa as representative of a distinct genus which he named Diatoma, and thus furnished the name that C. Agardh (1824) adopted for the group (as Diatomeae) and by which it is now commonly known. The most significant contributor to knowledge of this group during this early period was Nitzsch (1817). He gave the first useful illustrations of members of the class and also recognized their prismatic quality, which he considered a ma- jor character of the group. He carefully studied the multiplication of the rod- like forms by longitudinal division and pointed out, among other things, that the individuals did not lose their form after death. Nitzsch divided diatoms into two groups, animal and plant, according as they exhibited movement or not. Until 1832 members of this class were regarded partly as animals (the motile forms) and partly as algae (the nonmotile forms), although several botanists (C. Agardh, 1817, 1824, 1830-1832; Lyngbye, 1819; and others) had no hesi- tation in referring the entire group to the algae. In fact, C. Agardh in 1824 established for them the order Diatomeae, one of six which he recognized in the algae. Ehrenberg (1832, and many later publications), to the contrary, re- garded all diatoms as animals, without reservations, placing them in the family of rod animalcules (Bacillaria). C. Agardh, Ehrenberg, and others grouped the desmids with the diatoms. Kiitzing (1833b) was the first to recognize clearly the differences between these two groups of organisms, especially as regards the nature of the cell wall. Later, in a comprehensive monograph on the diatoms, Kiitzing (1844) elaborated on his earlier observations on the composition of the shell and pointed out that it is composed of silica. In this monograph Kiitzing also concerned himself with the classification of these organisms, and on the basis of the structure of the frus- tule, recognized a total of nineteen families (including one comprised largely of silicoflagellates). Thwaites (1847, 1848) was the first to observe the process of conjugation in diatoms. At first (1847) he did not comprehend the significance of his ob- servations but in 1848 he fully suspected that these phenomena were instances of a sexual process. In a monograph on diatoms published in 1853, Eabenhorst corrected cer- tain of Ehrenberg's and Klitzing's errors with respect to the structure of the frustule. In this publication, Eabenhorst considered the diatoms as constitut- ing an autonomous class of algae, which had no equal among living things as regards the sharpness of characters as shown by their peculiar type of shell. Previously, however, Harvey (1836) had regarded the diatoms (including the desmids) as forming one of the four divisions into which he divided the algae. Turpin in 1828 (b) expressed the view that the diatom shell consisted of three pieces, instead of two as had previously been believed, two valves, and a girdle. This view was adhered to until Wallich (1858, 1860b) pointed out that the girdle actually consisted of two connecting bands, one fitting over the other. From the point of view of the distribution of these pieces at cell division, MPfNFUSS: CLASSIFICATION OF THE ALGAE 153 Wallicli did not realize the significance of his discove^-. In 1869 Macdonald and Pfitzer independently of each other i)ointed out that, since at division a new valve and a new connecting band are formed within each of the two con- necting bands of the parent cell, one of the daughter cells is smaller than the other, which is of the same size as the parent. (That there occurs a decrease in the size of the frustule of a species at each cell division was suspected previ- ously by Griffith and Ilenfrey, 1856, p. 201.) Through continued division, cells are thus formed whose dimensions are api)reciably below the maximum charac- teristic of the species. Ultimately the cells would be too small to undergo further division and the race would perish unless a periodic reestablishment of maximal size occurred. Both Macdonald and Pfitzer considered the process of conjuga- tion as probably providing the required rejuvenescence. This postulate gained substance through the earlier observation of Braun (1851) that the cell de- veloping from a zygote is larger than the parent cells. Pfitzer also noted that in some instances a rejuvenating spore was produced by only one cell. Irrespec- tive of their method of formation, Pfitzer called these spores auxospores (en- larging spores). Two years later Pfitzer (1871) furnished abundant evidence favoring not only the concept of a reduction in the size of diatom cells through vegetative cell division but the reestablishment of the maximal size of the species through the process of auxospore formation. In this work he also gave the first detailed account of the living part of the diatom cell, the protoplast, which previously had received only slight attention, and introduced the characters presented by the plastids in the classification of these organisms. It will be recalled that largely because of their movement certain (or all) diatoms were for a long time regarded as animals. The exact method of move- ment of these organisms remained a matter of conjecture for more than a hundred years after the first species were described. In a series of papers start- ing in 1889 0. Miiller produced evidence that the movement of the cells is owing to cytoplasmic streaming along the raphe. Although the detailed mechanics of the process are not yet entirely understood, IMiiller's interpretation is still ac- cepted as the most plausible explanation of the phenomenon. In 1895 Miiller also published a valuable paper on the axial relations and planes of symmetry in diatoms, and coined, among others, the terms epitheca and hypotheca to de- note the larger and smaller halves, respectively, of the frustule. Klebahn in 1896, working on Rhopalodia gihha, a member of the order Pen- nales, was the first to obtain cytological results suggesting that diatoms are diploid and that meiosis occurs during gametogenesis. Further evidence of this was produced by Karsten in 1899, and in 1912 he gave convincing proof of the diploid nature of another member of the Pennales. Since that time various au- thors (Geitler, 1927a, 1927b, 1928; Cholnoky, 1928, 1933a; Meyer, 1929; Subrah- manyan, 1947) have confirmed the fact that the Pennales are diploid and that meiosis precedes auxospore formation. Since a conjugation of cells was not known to occur during auxospore formation in the Centrales, it was believed for a number of years (cf. Oltmanns, 1922a) that these forms, to the contrary, were haploid and that auxospore formation is an asexual process. The first per- son to show that the Centrales were likewise diploid and that here auxospore for- mation also is a sexual process (autogamy) was Persidsky (1929, 1935). His 154 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES observations have been confirmed by Cliolnolvv (1933b) and more particularly by Iyengar and Subralimanyan (1942, 194-1), Stosch (1951a), and Geitler (1952), It would seem therefore that the Centrales and Pennales are not as remote from each other as has been supposed. In 1897 G. Murray observed in certain marine members of the order Cen- trales rounded protoplasmic bodies which he interpreted as reproductive cells. Since then these so-called microspores have been observed by a number of inves- tigators in various marine as well as freshwater Centrales. In some instances the microspores are provided with two lateral and in others with two terminal flagella of equal length. Stosch (1951a) observed with certainty only one flagellum. It has been thought that these microspores are gametes but actual proof of this was not forthcoming until Stosch (1951a, 1951b) and Geitler (1952) showed that in some species they are actually male gametes. For a review of the litera- ture on the microspores reference should be made to the works of Karsten ( 1928, pp. 167-175), Fritsch (1935, pp. 633-637), Subralimanyan (1946), and Stosch (1951a). Utilizing a concept introduced into the classification of diatoms by Grunow in 1860, Kirchner (1878) and Schiitt (1896) divided these organisms into two groups, called Circulares and Bilaterales by Kirchner (p. 41) and Centricae and Pennatae by Schiitt, on the basis of the shape and sjonmetry relations of the valves. West (1904) elevated these two groups to the rank of order, ac- cepting Schiitt 's designations, and Karsten (1928) changed the names to Cen- trales and Pennales. Rabenhorst (1853) was the first to consider the diatoms as constituting an independent class of algae, which he (1864) named Diatomo- phyceae. The currently accepted name, Bacillariophyceae, was proposed by Fritsch (1935, p. 7). Engler and Gilg (1924, p. 13) and Karsten (1928) have elevated the group to the rank of phylum (Bacillariophyta) but in general phy- cologists have adhered to the interpretation of Pascher (1914, 1921), who, largely on the basis of the formation of endoplasmatic cysts in certain Centrales (first correctly interpreted by Schiitt in 1888) comparable to those of Chrysophyceae and Xanthophyceae, has related them to the latter two classes. The advantages of the present system of classification of the diatoms, which is based largely on the characters presented by the siliceous shell, is that it is applicable to the many fossil representatives (which are of considerable economic importance) as well as the living forms. The arrangement presented below is essentially that of Hustedt (1930). Class Bacillariophyceae Fritsch (1935, p. 7) Syn.: Class Diatomophyceae Rabenhorst (1S64, p. 2); Order Pyritophyceae Stizen- berger (1860, p. 23) ; Division Bacillariophyta Engler et Gilg (1924, p. 13) Order CENTRALES (Schiitt) West orth. mut. Karsten (1928, p. 201) Family Coscinodiscaceae Kiitzing orth. mut. De Toni (1890, p. 915) Syn.: Thaumatodiscaceae Cleve orth. mut. De Toni (1894, p. 1010); Melosiraceae Kiitzing orth. mut. De Toni (1890, p. 913); Xanthio- pyxidaceae (Petit) De Toni (1890, p. 914) ; Discaceae Schutt orth. mut. Karsten (1928, p, 201) Family Asterolampraceae H. L. Smith orth. mut. De Toni (1890, p. 919) Syn.: Heliopeltaceae H. L. Smith orth. mut. De Toni (1890, p. 918); Actinodiscaceae (Schutt) Hustedt (1930, p. 56) Family Eupodiscaceae Kutzing orth. mut. De Toni (1890, p. 916) PAPENFUSS: CLASSIFICATION OF THE ALGAE 155 Family Rhizosoleniaceae (Petit) De Toni (1890, p. 921) Syn.: Soleniaceae (Scluitt) Karsten (1928, p. 202) Family Chaetoceraceae H. L. Smith orth. mut. De Toni (1890, p. 920) Family Biddulphiaceae Kiitzing orth. mut. De Toni (1890, p. 910) Syn.: Isthmiaceae Cleve orth. mut. De Toni (1890, p. 913); Hemiauli- daceae Heiberg orth. mut. De Toni (1890, p. 912) Family Anaulaceae (Schiitt) Hustedt (1930, p. 56) P'amily Euodiaceae (Schiitt) Hustedt (1930, p. 56) Family Rutilariaceae Pantocsek orth. mut. De Toni (1894, p. 1020) Order PENNALES (Schiitt) West orth. mut. Karsten (1928, p. 202) Family Diatomaceae (S. F. Gray) Diimortier (1829, p. 77) Syn.: Fragilariaceae Kiitzing orth. mut. De Toni (1890, p. 905); Meridionaceae Kiitzing orth. mut. De Toni (1890, p. 904); Trachy- spheniaceae (Petit) De Toni (1890, p. 904) ; Plagiogrammaceae (Petit) De Toni (1890, p. 906); Licmophoraceae Kiitzing ortli. mut. De Toni (1890, p. 907); Striatellaceae Kiitzing orth. mut. De Toni (1890, p. 907); Entopylaceae Grunow orth. mut. De Toni (1890, p. 909); Tabellariaceae Kiitzing orth. mut. West (1904, p. 281) Family Eunotiaceae Kiitzing orth. mut. Rabenhorst (1853, pp. vii, 8, 15) Family Achnanthaceae Kiitzing orth. mut. De Toni (1890, p. 900) Syn.: Cocconeidaceae Kiitzing orth. mut. De Toni (1890, p. 899) Family Naviculaceae Kiitzing orth. mut. Rabenhorst (1853, pp. ix, 9, 36) Syn.: Cymbellaceae Kiitzing orth. mut. De Toni (1890, p. 898); Gomphonemaceae Kiitzing orth. mut. De Toni (1890, p. 899); Amphi- tropidaceae (Pfitzer) De Toni (1890, p. 898) ; Amphipleuraceae Grunow orth. mut. De Toni (1890, p. 902) ; Cocconemaceae West (1904, p. 298) Family Epithemiaceae Grunow orth. mut. De Toni (1892, p. 776) Family Nitzschiaceae Grunow orth. mut. De Toni (1890, p. 901) Syn.: Cylindrothecaceae (Kirchner) De Toni (1890, p. 902) Family Surirellaceae Kiitzing orth. mut. De Toni (1890, p. 903) Phylum Pyrrophycophyta Characterization : This phylum as now delimited embraces the single class Dino- phyceae which includes two somewhat dissimilar groups of largely unicellular organ- isms, the subclasses Desmophycidae (desmokonts) and Dinophycidae (dinoflagellates, peridinians). The Dinophycidae comprise forms which in the vegetative condition are (1) unicel- lular and biflagellate,'' (2) unicellular and amoeboid, (3) unicellular, nonmotile, and in the form of small gelatinous aggregates (palmelloid types), (4) unicellular and non- motile, with a firm cell wall (coccoid types), or (5) in the form of multicellular attached filaments. These diverse types share two prominent features which suggest that they constitute a related assemblage. Firstly, the flagellated species and the motile reproductive cells of the nonmotile forms exhibit what Graham (1951) calls a "dinoflagellate orientation." In them the two flagella are inserted near each other and laterally on what is known as the ventral side. One of the flagella is usually threadlike and projects backwards. At its proximal end it lies in a ventral, longitudinal groove, the sulcus. The other flagellum is ribbon-shaped and encircles the cell. It lies in a transverse or spiral groove, the girdle. The second feature common to these organisms is found in the structure of the nucleus: the chromatin is contained in threads which are distinctly beadlike and this character persists throughout karyokinesis. Less distinctive features which the Dinophycidae share with the Desmophycidae are: (1) the possession by the photosynthetic forms of a complex of pigments which give them a greenish tan or golden brown color (where examined [Strain, 1951, p. 253] 5. Polykrikos is colonial and each individual in the chainlike colony is furnished with two flagella. Whether or not any of the cells are separated by transvei'se walls is not clear from the literature. 156 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES the pigments have been found to consist of chlorophyll a, chlorophyll e, beta carotene, and four xanthophylls, three of which, as far as we know, are peculiar to the Dino- phyceae) ; and (2) the storage of food in the form of starch or oil. The cells of members of the Dinophycidae are either naked or are provided, in the forms referred to as armored dinoflagellates, with a cellulose wall, the theca. (The family Amphilothaceae comprises a small number of poorly known marine genera which possess an elaborate internal skeleton that may be silicified.) In some forms the cell is adorned with cellulosic horns (e.g., Ceratium) or saillike processes (e.g., Ornitliocercus) which aid in flotation. In the thecate, flagellated forms, the theca is made up of a series of articulated plates (except in members of the small family Ptychodiscaceae, in which it is homogeneous), the number and arrangement of which are important characters in classification. As regards method of nutrition, the Dinophycidae include both photosynthetic and heterotrophic forms (saprophytes, ecto- and endoparasites, and types with holozoic nutrition). The genera Polykrikos and Nematodinium possess nematocysts comparable to those occurring in coelenterates. The subclass Demosphycidae includes forms which are less specialized than those belonging to the Dinophycidae. The motile stages are biflagellate (with the flagella dissimilar or showing different movements) but do not show a dinoflagellate organiza- tion. In the vegetative condition the cells are provided with a cellulose wall (except in Desmoniastix which is naked) that consists of two valves joined by an antero-posterior suture or that splits into two valves along an antero-posterior plane when the protoplast is caused to swell. The sulcus and girdle are lacking and the two flagella are anterior in position. (The genus Desmocapsa is nonmotile in the vegetative condition and forms small palmelloid aggregates.) The Desmophycidae are placed in the Dinophyceae pri- marily on account of the structure of the nucleus. As far as known, the chromatin threads show the same moniliform condition as is characteristic of the Dinophycidae. The usual method of reproduction is by cell division, which in some forms is effected while the cell is motile, in others during an immobile phase. Cysts with a thick wall and abundant stored food are produced in a number of species, especially those inhabit- ing fresh water. The occurrence of sexual reproduction in the pyrrophycophytes has been established with certainty only in two species (Gross, 1934; Diwald, 1938). History: ". . . and all the waters that were in the river were turned to blood. And the fish that was in the river died; and the river stank, and the Egyptians could not drink of the water of the river; and there was blood throughout all the land of Egypt." (Exodus, vii. 20, 21.) Although the luminescent members of this group and those which, when present in large numbers, give a blood-red color to water have attracted the attention of man for centuries, the freshwater Ceratium hirundineUa and Peri- dinium cinctum are the forms to have been described first in a sufficiently pre- cise manner to be recognized by later workers. They were described by 0. F, Miiller in 1773 as Bursaria hirundinella and Vorticella cincta. These two species and a marine form which Miiller described later were subsequently redescribed and illustrated by him in 1786 in his Animalcula infusoria fluviatilia et marina . . . Following the publication of several papers in the proceedings of the Berlin Academy describing new genera and species and other observations upon Dino- phycidae, Ehrenberg in 1838 gave the first treatment of them as a coherent group in his famed work Die Infusionsthiere als vollkommene Organismen. Ehrenberg (1838, p. 249) placed them, along with certain organisms which he erroneously classified with them, in his twelfth family, the Peridinaea or "Kranzthierchen" (wreath animalcules), which formed the last family in his group "Polygastrica anentera," the gutless stomach animalcules. PAPENFUSS: CLASSIFICATION OF THE ALGAE 157 Ehrenberg observed the posteriorly directed flagellum but erred in usually figuring this end of the cell as the anterior end. lie also failed to understand the nature of the transverse flagellum, which he interpreted as a transverse band (or at times as two transverse bands) of cilia. That the longitudinal flagellum is directed posteriorly was first pointed out by Perty in 1852. He (1852) was also the first to call attention to the existence of naked representatives of the group. In Gymnodinium xiherriyyium Allman in 1855 first observed the peculiar structure of the luicleus in this phylum. Carter in 1858 confirmed an earlier observation by Allman (1855) to the effect that these organisms at times formed resting stages. Furthermore, he noted the division of the protoplast of resting stages to produce new individuals, which circumstance, apparently more than the fact that some of these beings possess chlorophyll, caused him to conclude that dinophycids were plants rather than animals. Carter (1858) was also the first to establish that the wall of dinophycids, at least as far as the resting stages were concerned, is composed of cellulose. On account of the alleged presence of a transverse band of cilia, Claparede and Lachmann (1858-1859) created a .separate order Cilioflagellata for the dino- phycids, and regarded them as a connecting link between the flagellates and the ciliates. To these authors goes the credit for first pointing out that the desmo- phycid genus Prorocentrum is related to the dinophycids instead of the crypto- monads where Ehrenberg (1838) had placed it. In 1872, Allmann expressed the view that the highly modified Noctiluca is allied to the dinophycids. This luminescent genus had for a long time been asso- ciated with the coelenterates and in 1873 Haeckel created for it the order Cystoflag- ellata, but subsequent work has shown that Allman 's conclusion was well founded. In a paper devoted largely to investigations on bacteria. Warming in 1876 briefly referred to his observations on dinophycids. He announced the occur- rence of cellulose in the wall of the motile stages of these organisms and thus extended the earlier findings of Carter (1858) that the wall of resting stages consists of cellulose. Warming also established that in these organisms food is stored in the form of starch and that some of them possess a pigment similar to that of diatoms. On the basis of these significant observations. Warming con- cluded, as Carter (1858) previously had on much less secure grounds, that the dinophycids were algae. A new era in the study of these organisms started with the publication in 1878 and 1883 of the first and second fascicles of the third part of Stein's Der Organismus der Infusionsthiere. Stein not only gave the best systematic treat- ment that had yet been presented of the group but illustrated them abundantlj^ and with considerable accuracy. Some of his figures still rank among the best that have been produced of the species in question. Stein regarded the Dino- phyceae as animals and placed them (Stein, 1883) as a suborder, "arthrodele Flagellaten" (articulated flagellates), in his order "Flagellaten." He divided the suborder into the five families Prorocentrinen, Noctiluciden, Peridiniden, Dinophysiden, and Cladopyxiden. During the period that Stein was studying these organisms, Bergh (1881) also published a treatise on them. He regarded them as constituting an order Cilioflagellaten which he divided into two families : the Adinida, which included 158 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES the girdleless and anteriorly biflagellate Prorocentrum, and the Dinifera, which received the forms with "dinoflagellate" structure. The latter family he divided into the three subfamilies Dinophysida, Peridinida, and Gymnodinida. The de- scriptive appellations Adinida and Dinifera introduced by Bergh have been em- ployed in one form or another in the classification of the Pyrrophycophyta down to the present. One of the most important advances in our knowledge of the structure of the dinophycid cell since the time of Ehrenberg was made by Klebs in 1883. He established that in freshwater forms the alleged transverse band of cilia actually is a single flagellum that lies in the transverse groove. A year later (Klebs, 1884) he established that this is true also of marine forms. A second significant contribution made by Klebs (1883) was concerned with the nucleus. He described the jointed structure of the chromatin threads and recognized the systematic value of this feature. It will be recalled that Allman in 1855 had already noted this condition (an observation which appears to have been over- looked by Klebs), but it is through the work of Klebs that this peculiarity was first brought into focus. Klebs in 1883 believed that the Dinophyceae were thallophytes but that they occupied a seemingly isolated position among them. In 1884 he was inclined to think that these organisms might be related to some of the other yellow flagellates. Confirmation of Klebs's observations, both with respect to the single trans- verse flagellum and the structure of the nucleus, came forth quickly through the work of Biitschli (1885). In Bronn's Klassen und Ordnungen des Thier- Reichs, Biitschli (1883-1887) also gave a comprehensive treatment of the Dino- phyceae, including an excellent review of the history of knowledge of the group. In consequence of the new information concerning the flagellation, Biitschli abandoned the name Cilioflagellata given to these organisms by Claparede and Lachmann and substituted the designation Dinoflagellata (whorled flagellates?) which has remained as the popular name of the assemblage. Recent observations by Deflandre (1934) indicate, ironically, that the transverse flagellum of Gleno- dinium uliginosum bears a single row of cilia. The first formal recognition of the dinoflagellates as a group of plants came in 1890 and 1892 when AVarming (1890)*^ and Engler (1892) accepted them as a subdivision of the thallophytes. They were henceforth always included in treatises on the algae or the plant kingdom as a whole. An outstanding monograph on the structure of the cell in marine dinoflagel- lates was published bj^ Schiitt in 1895 as part of the results of the Plankton Expedition, and in 1896 the same author presented an excellent systematic treat- ment of the group, with the exception of certain forms such as Noctiluca and Polykrikos which were excluded. Schiitt (1896) divided the group into three families: (1) the Prorocentraceae, which included the terminally biflagellate forms, (2) the Gymnodiniaceae, which received the athecate forms with dino- flagellate organization, and (3) the Peridiniaceae, in which he placed the thecate forms with dinoflagellate organization. 6. According to Warming (1890, p. V) the dinoflagellates were first accepted as algae by Petersen and himself in 1889 in their Grundtrak af Forelcisninger over systematisk botanik for medicinske og farmaceutiske studerende. PAPENFUSS: CLASSIFICATION OF THE ALGAE 159 With few exceptions (e.g., Pyrocystis) the Pyrrophycophyta which had be- come known to science previous to 1912 were flagellated forms. In that year Klebs published his significant discovery of several nonmotile unicellular organ- isms that at certain stages in their development clearly revealed their relation- ship to the dinoflagellates. Two years later Pascher (1914) not only announced the discovery of a num- ber of additional nonmotile types but proposed a far-reaching revision of the classification of the dinoflagellates. He erected a phylum Pyrropliyta and ac- credited it with the three groups Cryptophyceae", Desmokontae, and Dinophy- ceae. The Dinophyceae received, in addition to the characteristic flagellated forms with dinoflagellate organization, those nonmotile genera with Gymnodi- nmm-like swarmers that had been discovered by Klebs and himself. Some of these forms had a palmelloid organization (his Dinocapsales), others had a coc- coid organization (his Dinococcales), and the single representative of a third group had a filamentous organization (his Dinotrichales). The Desmokontae included the forms which lacked a dinoflagellate organiza- tion throughout their life history. They were divided into the two orders Des- momonadales and Desmocapsales, the Desmomonadales receiving the four fami- lies Desmomonadaceae, Exuviaellaceae {nomen nudum), Prorocentraceae, and Dinophysiaeeae and the Desmocapsales accommodating the monogeneric family Desmocapsaceae. Knowledge of the pyrrophyeophytes has progressed by great strides during the past forty years. Space permits the consideration of but a few of the many investigations that have contributed to this advancement. For our knowledge of the parasitic dinoflagellates we are especially indebted to Chatton, who in 1920 published an extensive monograph on the morphology and taxonomy of these forms. Some of them are ectoparasites, others are endo- parasites, mostly on marine metazoa. Although these species bear little resem- blance to ordinary dinophycids, their relationship to them is clearly revealed by the structure of the motile reproductive cells. In a long series of publications, Kofoid and his associates contributed signifi- cantly to our knowledge of the motile dinophycids. In 1921 Kofoid and Swezy produced a monograph on the unarmored forms, based mostly upon their obser- vations of living material obtained in the vicinity of La Jolla, California. They described a number of new families and genera and presented a revision of the classification of Dinophyceae. Kofoid and Swezy did not follow Pascher (whose paper of 1914 they did not refer to) in the separation of the terminally bi- flagellate forms into a separate group, the Desmokontae. They regarded the Dinoflagellata as a subclass of the class Flagellata in the phylum Protozoa and treated the terminally biflagellate forms as an order Adiniferidea of this sub- class. The forms with "dinoflagellate" motile cells they placed in an order Diniferidea. With few exceptions, the genera which Pascher placed in his orders Dino- capsales, Dinococcales, and Dinotrichales were not considered by Kofoid and Swezy. In fact they regarded (p. 109) the genera described by Klebs (1912), namely, Phytodinium, Tetradinium, Stylodinium, and Gloeodinium, as more nearly related to the green algae than to the dinoflagellates. 7. The Cryptophyceae are now excluded from the phylum. 160 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCE^ In 1928 Kofoid and Skogsberg published an extensive monograph upon the dinoflagellates of the "Albatross" expedition. Three new families and five new genera were described in this work. The most extensive systematic treatises on the pyrrophycophytes are those of Lindemann, published in 1928 as a volume in the second edition of Engler and Prantl's Pflanzenfamilicn, and of Schiller, published in two volumes be- tween 1931 and 1933, and 1935 and 1937. Many of the currently accepted fami- lies were established by Lindemann. The volumes by Schiller appeared as part of the second edition of Rabenhorst's Kryptogamen-Flora von Deutschland, Os- terreich und der Schweiz but their scope is much more comprehensive than the title of the series suggests inasmuch as they treat of all the known living species. Valuable general accounts of the phylum have recently been given by Fritsch (1935) and Graham (1951). The classification presented in the synopsis below is a synthesis of the systems of Pascher, Lindemann, Schiller, Fritsch, and Graham. This arrangement de- parts in certain major respects from Pascher 's system and in conclusion it is deemed desirable to review briefly the more significant points in the evolution of this classification. It will be recalled that Pascher (1914, 1927a, 1931) accredited the Pyrro- phycophyta with the three groups Desmokontae, Cryptophyceae, and Dinophy- ceae. Fritsch (1935) treats the Cryptophyceae as a distinct class, which at best may be only distantly related to the other two groups. Graham (1951) has further emphasized the distinctness of the Cryptophyceae, especially as regards the structure of the nucleus, and in agreement with him they are here consid- ered as a separate class appended to the Pyrrophycophyta. Fritsch (1935) in agreement with many earlier workers, regards the Desmo- kontae and the Dinophyceae of Pascher as more closely related than is implied by Pascher's system and treats them as groups, Desmokontae and Dinokontae, belonging to a common class, the Dinophyceae. Pascher (1914) accredited the Desmokontae with the two orders Desmomona- dales and Desmocapsales. Fritsch (1935, p. 672) produces convincing reasons for placing the single genus Besmocapsa, upon which Pascher based the Desmo- capsales, in the family Desmomonadaceae of the order Desmomonadales. In the order Desmomonadales Pascher (1914) had placed four families, viz., Desmomonadaceae, Exuviaellaceae, Prorocentraceae, and Dinophysiaceae. In 1928 Lindemann established the order Thecatales^ for the Prorocentraceae, leav- ing the order in the Desmokontae (or Adiniferae as he called this group). Pas- cher (1931), Schiller (1931), Fritsch (1935), and Graham (1951) have accepted this order, except that Pascher and Graham call it Prorocentrales. Lindemann also established an order Dinophysiales for the Dinophysiaceae, added to it a second family, the Amphisoleniaceae, and removed the order to the Dinokontae (or Diniferae as he called this group). Pascher (1931), Schiller (1931), who enriched the order with two more families, and Fritsch (1935) ac- cept the order Dinophysiales but retain these forms in tlie Desmokontae. Graham (1951, p. Ill), however, produces well-founded arguments for referring the Dinophysiales to the Dinokontae, with which group Kofoid and Skogsberg (1928) as well as Lindemann had related it. 8. As "Klasse," but Liudemann's classes are actually all orders. PAPENFUSS: CLASSIFICATION OF THE ALGAE 161 Present-day classification of the flagellated Dinophycidae is largely based on the systems of Kofoid, Lindemann, and Schiller. Whereas the Dinophycidae were represented by two families in I he arrangement of Schiitt, published in 1896, they arc here segregated into eight orders and a total of 38 families. Phylum PYRROPHYCOPHYTA Papenfuss (1946, p. 218) Syn.: Pyrrophyta Pascher (1914, p. 153) Class DiNOPHYCEAE Fritsch (1935, pp. 8, 665) Subclass DF:sMOPnY('iDAE (Pascher) Graham orth. mut. Papenfuss Syn.: Desmokontae Pascher (1914, p. 149, as "Reihe"); Subdivision Adiniferae (Bergh) Lindemann (1928, p. 36); Order Adiniferidea (Bergh) Kofoid et Swezy (1921, p. 108) ; Subclass Desmokontae (Pascher) Graham (1951, p. 105) Order DESMOMONADALES Pascher (1914, p. 148) Syn.: Desmocapsales Pascher (1914, p. 149); Athecatales Lindemann (1928, p. 36) Family Desmomonadaceae Pascher (1914, p. 149) Syn.: Desmocapsaceae Pascher (1914, p. 149; cf. Fritsch, 1935, p. 672); Haplodiniaceae Lindemann (1928, p. 36) ?Family Adinimonadaceae Schiller (1931, p. 9) Order THECATALES Lindemann (1928, p. 37) Syn.: Tribe Thecatoidae Kofoid et Swezy (1921, p. 106), nomen nudum; Pro- rocentrales Pascher ex Graham (1951, p. 114) Family Prorocentraceae Engler (1892, p. 6) Syn.: Exuviaellaceae Pascher (1914, p. 148), nomen nudum Subclass DINOPHYCIDAE (Fritsch) Graham orth. mut. Papenfuss Syn.: "Group" Dinokontae Fritsch (1935, pp. 670, 679); "Reihe" Dinophyceae Pascher (1914, p. 151); Subclass Dinokontae Graham (1951, p. 105) Order GYMNODINIALES (Poche) Lindemann (1928, p. 39) Syn.: Amphilothales (Kofoid et Swezy) Lindemann (1928, p. 68; cf. Zimmer- mann, 1930, pp. 438-440; Schiller, 1935, p. 1) Family Pronoctilucaceae Lebour orth. mut. Lindemann (1928, p. 39) Family Gymnodiniaceae (Bergh) Schiitt (1896, pp. 1, 2) Family Polykrikaceae Kofoid et Swezy orth. mut. Lindemann (1928, p. 46) Family Noctilucaceae Kent orth. mut. Lindemann (1928, p. 47) Family Warnowiaceae Lindemann (1928, p. 51) Family Amphilothaceae Kofoid orth. mut. Lindemann (1928, p. 68) Syn.: Gymnasteraceae Poche orth. mut. Lindemann (1928, p. 69); Gymnosclerotaceae Schiller (1935, p. 1) Order BLASTODINIALES Schiller (1935, p. 8) Family Paradiniaceae Schiller (1935, p. 15) Family Blastodiniaceae Chatton orth. mut. West (1916, p. 50) Family Syndiniaceae Chatton orth. mut. Schiller (1935, p. 53) Family Endodiniaceae Schiller (1935, p. 61) Family Ellobiopsidaceae Schiller (1935, p. 62) Order DINOPHYSIALES (Kofoid) Lindemann (1928, p. 72) Family Dinophysiaceae (Bergh) Engler orth. mut. Pascher (1914, p. 158) Family Amphisoleniaceae Lindemann (1928, p. 77) Family Ornithocercaceae Kofoid et Skogsberg orth. mut. Schiller (1931, p. 192) Family Citharistaceae Kofoid et Skogsberg orth. mut. Schiller (1931, p. 255) Order PERIDINIALES Schutt (1896, p. 1) Syn.: Kolkwitziellales Lindemann (1928, p. 70) Family Ptychodiscaceae (Schutt) Lemmermann (1899, p. 362) Syn.: Kolkwitziellaceae Lindemann (1928, p. 71; cf. Schiller, 1935, p. 75) 162 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Family Pyrophacaceae Lindemann (1928, p. 96) Syn.: Glenodiniopsidaceae Schiller (1935, p. 80) Family Glenodiniaceae (Scliiitt) Lemmermann (1899, p. 361) Syn.: Kyrtodiniaceae Schilling (1913, p. 12); Dinosphaeraceae Linde- mann (1928, p. 84; cf. Schiller, 1935, p. 99) Family Peridiniaceae Ehrenberg orth. mut. Engler (1892, p. 6) Syn.: Krossodiniaceae Schilling (1913, p. 30) Family Goniaulaceae Lindemann (1928, p. 84) Family Congruentidiaceae Schiller (1935, p. 320) Family Protoceratiaceae Lindemann (1928, p. 83) Family Ceratiaceae (Schiitt) Lindemann (1928, p. 91) Syn.: Heterodiniaceae Lindemann (1928, p. 95; cf. Schiller, 1937, pp. 327-432) Family Goniodomaceae Lindemann (1928, p. 94) Family Ceratocoryaceae (Schiitt) Lindemann (1928, p. 98) Family Oxytoxaceae (Schiitt) Lindemann (1928, p. 97) Family Cladopyxiaceae (Kofoid) Poche orth. mut. Lindemann (1928, p. 99) Family Ostreopsiaceae Lindemann (1928, p. 96) Family Podolampaceae (Schiitt) Lindemann (1928, p. 100) Family Lissodiniaceae Schiller (1937, p. 480) Order RHIZODINIALES Pascher (1931, pp. 320, 326) Family Amoebodiniaceae Pascher (1931, p. 326) Syn.: Dinamoebaceae Pascher (1916b, p. 135) Order DINOCAPSALES Pascher (1914, p. 151) Family Gloeodiniaceae Pascher ex Schiller (1937, p. 482) Syn.: Dinocapsaceae Pascher (1914, p. 158), nomen nudum Order DINOCOCCALES Pascher (1914, p. 151) Family Hypnodiniaceae Pascher (1931, p. 326), nomen nudum Family Phytodiniaceae Klebs (1912, p. 443) Syn.: ?Pyrocystaceae (Schiitt) Poche orth. mut. West (1916, p. 55); Dissodiniaceae Graham (1951, p. 116), nomen nudum Family Protaspidaceae Skuja (1939b, p. 116; cf. Skuja, 1948, p. 375) Family Stylodiniaceae Pascher (1931, p. 326), nomen nudum Order DINOTRICHALES Pascher (1914, p. 151) Family Dinotrichaceae Pascher (1914, pp. 151, 158, 160; 1927a, pp. 2-15; 1931, p. 326) Family Dinocloniaceae Pascher (1927a, p. 15; 1931, p. 326) Classes of Uncertain Systematic Position CLASS CRYPTOPHYCEAE Characterization: This class embraces less than two dozen genera of highly special- ized, asymmetrical, compressed, usually flagellated, pigmented or rarely colorless, uni- cellular organisms. The cells have a firm periplast but lack a wall. The flagella, of which there are two, are of slightly unequal length and are somewhat ribbon-shaped with a tapering end. They are usually inserted terminally but are lateral in a few forms. A few species are palmelloid and at least one monotypic genus {Tetragonidium) is coccoid in organization. The majority of the pigmented forms are provided with two parietal plastids of a brown, red, blue, green or bluegreen color. In rare and somewhat doubtful instances (cf. Pringsheim, 1944, p. 148) the cells appear to contain several discoid plastids. Pyrenoids and an eye-spot may or may not be present. The colorless forms are sapro- phytic or holozoic. In the motile genera and in the zoospores of the immobile forms there is a superficial curved furrow which extends backward from the place of insertion of the flagella. In many genera a "gullet" extends into the protoplast from the point of insertion of the flagella. The "gullet" may or may not be lined on the side adjacent to the protoplast with trichocysts, or (in the Cryptochrysidaceae) the trichocysts may be situated in the PAPENFUSS: CLASSIFICATION OF THE ALGAE "163 furrow. The cells usually contain one contractile vacuole. Reserve food is deposited as starch or starchlike compounds. Many of the Zooxanthellae growing symbiotically in the tissues of radiolarians and corals are members of this class. History: Knowledge of the Cryptophyceae begins with the year 1832, when Ehrenberg described Cryptomonas and Chilomonas. In 1838 he erected for Cryj)- tomonas (and certain other forms which have since been shown to belong else- where) tlie family Crj'ptomonadina, and referred it to his group 'Tolygastrica anentera." Dujardin (1841, p. 270) placed the cryptomonads along with a number of other flagellated organisms in his order "Infusoires pourvus d'un ou plusieurs filaments flagelliformes servant d'organes locomoteurs. — Sans bouche," for which group Cohn (1853) later proposed the designation Flagellata. The cryptomonads retained their position in the Flagellata for a long time (Stein, 1878; Blitschli, 1883-1887; Klebs, 1892; Senn, 1900; Lemmermann, 1907- 1910). Various early authors (e.g., Cienkowsky, 1870; Schmitz, 1882; Dan- geard, 1889) regarded them as algae, but general acceptance of them as a group of plants begins with the year 1900 when Senn gave a treatment of them in Engler and Prantl's Natilrlichen Pflanzenfamilien. Xlebs (1892, p. 392) circumscribed the group Flagellata in such a way that it included only the five subgroups Protomastigina, Polymastigina, Euglenoi- dina, Chloromonadina, and Chromomonadina. The Chromomonadina comprised, according to his system, the two families Chrysomonadina Stein and Crypto- monadina Ehrenberg. Although he placed these two families of essentially yellow-brown organisms in a common group, Klebs emphasized that the crypto- monads stood well apart from the chrysomonads. He especially drew attention to the fact (p. 420) that the cryptomonads stored starch, which was not known to occur in other Flagellata (according to his circumscription of this assem- blage), and in this respect agreed with the dinoflagellates. Klebs regarded the Flagellata as standing intermediate between plants and animals and believed they were the progenitors of various other lower organ- isms. He was impressed by the prominent plantlike features of many members of his group Chromomonadina and said (p. 278) that one could refer to them as chrysophytes, a designation that was later formally adopted by Pascher (1914) as the phyletic name for the chrysomonads, heterokonts, and diatoms. In agreement with Klebs (1892), Pascher in 1911 (b) believed in an alliance between the cryptomonads and the chrysomonads, but he also pointed to the possibility of a relationship between the cryptomonads and the dinoflagellates, a view which had been held previously by Bergh (1881) and Blitschli (1883- 1887, 1885). In fact, Ehrenberg in his day had placed Prorocentrum in his family Cryptomonadina. Pascher finally in 1914 removed the cryptomonads from the vicinity of the chrysomonads and placed them as a group, Cryptophy- ceae, along with the Desmophycidae and the Dinophycidae in the newly erected phylum Pyrrophyta. Fritsch (1935), Smith (1938), Pringsheim (1944), Graham (1951), and others have accepted the class Cryptophyceae but various authors, especially Pringsheim and Graham, have been skeptical of its presumed relationship with the Dinophyceae. Graham has in fact removed the class from the Pyrrophyco- phyta — primarily on the basis of the difference in nuclear structure. 154 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES The following arrangement is a synthesis of the systems of Pascher (1931) and Pringsheim (1944). Pringsheim has pointed to certain weaknesses in the classification of Pascher and more recently Skuja (1948) has added to the class the new family Senniaceae and has removed the Nephroselmidaceae to the Vol- voeales in the green algae. Class Cryptophyceae (Pascher) Fritsch, in West (1927, p. 387) Non Cryptophyceae Thuret, m Le Jolis (1863, pp. 13, 25), nomen nudum Order CRYPTOMONADALES Senn, in Engler (1903, p. 7) Syn.: Phaeochrysidales Pascher (1910, p. 9); Phaeocapsales Pascher (1912a, p. 196); Cryptocapsales Pascher (1931, p. 325) Family Cryptochrysidaceae (Pascher) Pascher (1931, p. 325) Family Cryptomonadaceae Ehrenberg orth. mut. Senn, in Engler (1903, p. 7) Syn.: Phaeocapsaceae De Ton! (1895, p. 591) ; Phaeoplakaceae Pascher (1931, p. 325); Chiiomonadaceae Lemmermaun (1908, p. 473) Family Cyathomonadaceae Pringsheim (1944, p. 149) Family Kathablepharidaceae Skuja (1939b, p. 96) Family Senniaceae Skuja (1948, p. 367) Order CRYPTOCOCCALES Pascher (1914, p. 150) Family Cryptococcaceae Pascher (1931, p. 325) See the figures of Tetragonidium by Thompson in Smith (1950, p. 636). CLASS CHLOROMONADOPHYCEAE Characterization: This class embraces a few highly specialized unicellular, anteriorly biflagellate genera (excepting Monomastix, which is uniflagellate, and Mer atrichia, which is laterally biflagellate). The cells are naked, provided with a delicate periplast, meta- bolic, flattened, dorsiventral, ovoid or pear-shaped, and usually possess a longitudinal groove on the ventral surface. The flagella issue from a slight depression — one is directed forwards and the other trails behind along the ventral surface. They are of the same length, except in Thaumatomastix and Yacuolaria viridis, in v/hich the trailing flagellum is longer than the other, and in Gonyostomum, in which it may be shorter or longer than the other. The majority of the forms are green and are provided with numerous discoid chromatophores containing a preponderance of xanthophylls. Nothing is known about the composition of the pigment complex. Two of the genera {Reckertia, Thaumatomastix) are colorless and presumably holozoic. Food is stored as oil. An eye-spot is lacking. Contractile vacuoles are present. Some forms (e.g., Trentonia, Gonyostomum) are pro- vided with an anterior cavity connected by a duct to the exterior and some (e. g., Mero- trichia, Gonyostomum) possess trichocysts. Reproduction is by longitudinal division of the cell. Cysts with a firm gelatinous wall may be produced. History: This small class of only seven genera was first established as an autonomous group (Chloromonadina) by Klebs (1892, pp. 292, 391-394) who referred to it the genus Yacuolaria Cienkowski (1870) and forms belonging to Gonyostomum Diesing (1866) and Merotricliia Mereschkowsky (1879) as cur- rently delimited. The first species to have been described sufficiently well to be recognized by later workers is Gonyostomum semen which was described by Ehrenberg (1853) as Monasf semen. Blitschli (1884, p. 819) placed the members known at the time of his writing together with a variety of unrelated genera in his family Coelomonadina. Formal recognition of the Chloromonadina as a group of plants begins with Engler (1898, p. 8) and Luther (1899, p. 19) who independently erected for them an order Chloromonadales. Luther placed the order in his class Heterokontae. f>APENFUSS: CLASSIFICATION OF THE ALGAE 155 Senn (1900, pp. 170-173) maintained the chloromonads as a separate group in the Flagellata and gave a systematic treatment of the complex. Some of the genera wliich he referred to the group have since been shown to belong else- where. Since the time of Senn, Lemmermann (1907-1910), Pascher (1913c), Skuja (1948) and Huber-Pestalozzi (1950) have given systematic treatments of various genera comprising the complex, and Drouet and Cohen (1935, 1937) have given a good account of the morphology of Gonyostomum semen. Fritsch {in West, 1927, p. 405) elevated the group to the rank of class. The majority of authors have regarded the Chloromonadophyccae as an iso- lated group of flagellates of uncertain relationship (cf. Fritsch, 1935, p. 723; Smith, 1950, p. 625). Prescott (1951, p. 421) has in fact erected a phylum Chloromonadophyta for the group. Oltmanns (1922a, p. 44) recognized the cor- respondence between the chloromonads on the one hand and the euglenids and the cryptomonads on the other, but made it clear that he like many others was not sure that this implied a definite relationship. Both Skuja (1948) and Huber-Pestalozzi (1950, p. 2) place the Chloromo- nadophyccae, Cryptophyceae, and Dinophyceae as classes in the phylum Pyrro- phycophyta. The uniflagellate genus Monomastix shows relationships to both the Chloromonadophyccae and the Cryptophyceae. Huber-Pestalozzi (1950, p. 2) considers it the type of a subclass in the Cryptophyceae whereas Skuja (1948, p. 344) places it in the Chloromonadaceae. Although neither the Chloromonadophj^ceae nor the Cryptophyceae appear to be closely related to the Dinophyceae (see the section on the Cryptophyceae regarding Pringsheim's [1944] and Graham's [1951] doubts about the presumed relationship between the Cryptophyceae and Dinophyceae) it is not inconceiv- able that the Chloromonadophyccae and the Cryptophyceae are at least dis- tantly allied and they are therefore here placed near each other as classes ap- pended to the Pyrrophycophyta. Outstanding points of agreement between these two classes are: (1) the cells are naked and more or less dorsiventral; (2) a longitudinal furrow is present in the cells of both classes; (3) the majority of the forms in both groups are anteriorly bifiagellate; (4) trichocysts are present in certain members of both groups; (5) some chloromonads have a cavity at the anterior end of the cell (connected to the exterior by a duct) which is com- parable to the "gullet" of some cryptomonads. A conspicuous but phylogenetically perhaps insignificant difference between the two groups is the storage of reserve food as starch or starchlike compounds in the Cryptophyceae and as oil in the Chloromonadophyccae. The flagella in the two groups are also of a somewhat different structure and are arranged differently. The systematic arrangement here adopted is essentially that of Huber- Pestalozzi (1950), except that Moyiomastix is considered, in agreement with Skuja (1948), as belonging to the Chloromonadophyccae instead of the Cryptophyceae. Class Chloromonadophyceae (Klebs) Fritsch orth. mut. Drouet et Cohen (1935, p. 423) Syn.: Raphidophycinees Chadefaud (1950a. p. 789) Order MONOMASTIGALES (Huber-Pestalozzi) Papenfuss, stat. nov. Syn.: Subclass Monomastiginae Huber-Pestalozzi (1950, p. 2) Family Monomastigaceae Huber-Pestalozzi (1950, p. 2) 166 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Order CHLOROMONADALES Engler (1898, p. 8, as "Reihe") Syn.: Raphidomonadales Chadefaud (1950a, p. 789) Family Vacuolariaceae Luther (1899, p. 19) Syn.: Chloromonadaceae Engler (1898, p. 8, nomen nudum) ex Prescott (1951, p. 421); Gonyostomaceae Lemmermann (1907-1910, p. 478); Tliaumatomastigaceae Skuja (1939b, p. 99); Thaumatonematidae Poche (1913, p. 155) Phylum Phaeophycophyta Characterization: The algae belonging to this phylum owe their characteristic olive- green to dark brown color to the presence in their plastids of certain xanthophylls, espe- cially fucoxanthin (which is peculiar to them and to diatoms), that mask the other pig- ments: chlorophyll a, chlorophyll c, and beta-carotene. Depending upon the genus or species, the cells possess one to many plastids of varying form and size. Pyrenoids have been recorded for a number of species but these structures may not be true pyrenoids. Ordinarily the cells are uninucleate. The cell wall is differen- tiated into an inner cellulosic and an outer pectic portion consisting usually of a gumlike substance, algin, which has many economic uses. Calcification of the wall occurs in the genus Padina. The known food reserves are the polysaccharide laminarin, the alcohol mannitol, and fats. The simplest Phaeophycophyta, as exemplified by certain members of the order Ectocarpales, have a branched, uniseriate, filamentous, and frequently microscopic thallus. The orders Laminariales and Fucales are comprised of morphologically elaborate forms that in size, degree of external differentiation, and complexity of structure surpass all other algae. Growth in length of the thallus is apical or marginal as the result of a single initial or a row of initials. Some show diffuse growth. Many possess an intercalary meristem. In the Laminariales it is situated between the stipe and blade and contributes cells to both; in other groups (e. g., Desmarestiales) it is located at the base of a terminal hair. Growth in width, thickness, or girth of the thallus is effected by repeated longitudinal division of the first-formed segments, or by the formation of radially directed filaments. In many forms the surface layer of cells remains meristematic and through periclinal division contributes to the growth in girth or thickness. The majority of brown algae show an alternation of generations. The two genera- tions may be morphologically identical (isomorphic) or dissimilar (heteromorphic). The diploid asexual generation forms either unilocvilar sporangia, plurilocular sporangia, or both. The unilocular sporangium develops from a single cell, which is not partitioned by walls. It is the seat of meiosis. The haploid zoospores (aplanospores in the Dictyotales) that are produced in it give rise to sexual plants. The plurilocular sporangia are formed by a linear series of cells (or rarely a single cell) that are divided into compartments, in each of which is formed a single zoospore. No reduction of chromosome number occurs in these sporangia. Their zooids give rise to other diploid, sporophytic plants. The sexual plants are monoecious or dioecious and they are either isogamous, anisog- amous, or oogamous. In isogamous and anisogamous forms the gametangia are pluriloc- ular organs that in structure agree with the plurilocular sporangia of the sporophytic generation. In oogamous species one or more eggs are produced in each oogonium and one or many sperms in each antheridium. The motile reproductive cells are pear-shaped, usually possess an eye-spot, and are laterally biflagellate." The anteriorly directed flagellum is longer than the posteriorly directed one, except in the Fucales, whose sperms have a long posterior and a short anterior flagellum. Longest (1946) has shown that in Ectocarpus the long anterior flagellum is of the tinsel type, an observation that has been confirmed by Manton and 9. This is true even of the sperms of Dictyota, which were described by Williams (1904b) as possessing only an anterior fiagellum. Shortly afterwards, however, he dis- covered the presence also of a short posterior flagellum, a fact recently mentioned in his obituary notice by Knight (1947). In a paper that appeared after the manuscript of this chapter had gone to press, Manton, Clarke, and Greenwood (1953), on the contrary describe and illustrate the sperms as being uniflagellate. PAPENFUSS: CLASSIFICATION OF THE ALGAE 167 Clarke (1951a) with respect to Pylaiella and Laminaria. It is of interest to note that in the sperms of Fucus it is also the anterior flagellum that is of the tinsel type (Manton and Clarke, 1951b). History: For some fifty years after the publication of Linnaeus' Species pluntarum (1753) almost all nonmembranous parenchymatous or pseudoparen- chjTiiatous algae (brown, red, and green forms such as Caulerpa) were referred to the genus Fucus. (For the long pre-Linncan history of this genus the reader is referred to tlie interesting article by Church, 1919a.) Stackhouse in his Ne- reis hritannica (1795-1801) and later in his Tentamen marino-cryptogamicimi (1809) was the first to recognize the heterogeneity of this genus which he ac- cordingly subdivided into a large number of genera, a few of which {Chorda, AscophyUum, Bifurcaria) arc still accepted as genera of brown algae. Largely on the basis of their brown color, Lamouroux (1813) erected a group ("ordre"), Fucacees, for some of the genera of this phylum. He, however, ex- cluded from the Fucacees the members of the Dictyotaceae, which he regarded as representative of a separate ''ordre," Dictyotees. C. Agardli (1817) changed Lamouroux' designation to Fucoideae and consid- ered these algae as constituting one of the five sections into which he divided the algae. Like Lamouroux, C. Agardh failed to make a sharp separation of the algae on the basis of color. In 1817, he placed the Dictyotees of Lamouroux in his section Ulvoideae. In 1824 he removed them to the Fucoideae but he still kept the filamentous brown algae in the Confervoideae. With few exceptions, the autonomy of the brown algae was henceforth ac- cepted as an established fact. On account of their brown pigment, Harvey (1836) named them Melanospermeae, which designation was changed to ]\Iel- anophyceae by Ruprecht (1851). Thuret (1850) created the name Phaeosporeae for one of the major taxa into which he (1855) divided the group. De Bary (1881) coined the designation Phaeophyceae, which is now generally accepted as the class name of the group. The discoveries relating to sexuality and of alternation of generations in the Phaeophyceae contributed immensely to an understanding of the life histories of thallophytes. The two kinds of reproductive organs characteristic of a large majority of these algae were named oosporangia and trichosporangia by Thuret in 1850 (pp. 235, 236), but shortly afterwards (1855, p. 15) he proposed the subsequently employed terms unilocular and plurilocular sporangia. Thuret found that the swarmers from the two kinds of sporangia were morphologically similar, except for size, and remarked (1850, p. 236), "J'ai vu d'ailleurs germer les uns et les autres, ce qui prouve suf!isamment leur complete identite." Studying Fucus, Thuret in 1853 observed for the first time in brown algae that only eggs to which sperms had had access would germinate. His classical illustrations of the reproductive organs were published in 1854. Strasburger in 1897 saw the fusion of the egg and sperm nuclei and established that the plants are diploid. His cytological observations were confirmed by Farmer and Wil- liams (1896, 1898) and by Yamanouchi (1909a), who also esta])lished that meio- sis occurs during the first two divisions of the primary nucleus of the oogonium and antheridium. Fucus (and this is true of related genera also) thus was shown to have a life history analogous to that of animals. The next brown alga in which a conjugation of gametes was observed is 168 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Zanardinia. Reinke (1877, 1878) found that in this genus the swarmers from the plurilocular organs (of which there are two kinds — some with large and some with small locules — both borne on the same plant) are gametes which con- jugate in pairs, the smaller zooids functioning as male gametes. This was the first observation of the actual fusion of gametes in brown algae. Eeinke noted that in Zanardinia the unilocular sporangia occurred on separate plants and found that the swarmers from these sporangia always germinated directly. In view of the occurrence of an alternation of generations in higher cryptogams (as had become well established by this time through the pioneering studies of Hof- meister and others), Reinke had no hesitation in interpreting his observations as indicating the occurrence in Zanardinia of a similar alternation between gametophytic and sporophytic generations. This, then, is the first brown alga which was considered as showing this phe- nomenon. At first botanists hesitated to accept Reinke's interpretation but its accuracy was established cytologically by Yamanouchi (1911, 1913). Reinke (1878) and Falkenberg (1879) also observed the fusion of the zooids from the plurilocular organs of Cutleria, a genus closely related to Zanardinia. Both of them were of the opinion that another alga known as AgJaozonia (which bears only unilocular organs) represented the sporophytic generation of Cut- leria. The evidence in favor of this view accumulated in the course of the next few decades and finally Yamanouchi (1909b, 1912) furnished cytological proof of it. Berthold (1881b) studying the classical Ectocarpus siliculosus at Naples found that also in this genus the zooids from the plurilocular organs were gametes. He also saw the fusion of the gamete nuclei, an observation which had been made only once before in plants — by Schmitz (1879c) in Spirogyra. Ber- thold was unable to determine the role of the unilocular sporangia of Ectocarpus since none of the plants obtained in the sea at Naples bore any. In consequence of the observations of Reinke, Falkenberg, and Berthold re- garding the gametic role of the zooids from the plurilocular organs of Zanar- dinia, Cutleria, and Ectocarpus a firm conviction developed among botanists (and was adhered to for almost half a century) that the plurilocular organs of brown algae were always gametangia and the unilocular organs sporangia. Not infrequently it was found (e.g., by Berthold, 1881b; Sauvageau, 1896a, 1896b, 1897; Oltmanns, 1899; Kuckuck, 1891) that the zooids from the plurilocular organs did not conjugate but germinated directly. To explain this asexual be- havior the theory was usually advanced that the gametes had lost their sexual power and germinated parthenogenetically. That this explanation was incorrect was shown by Knight (1923, 1929). Study- ing Pylaiella (a genus related to Ectocarpus) and Ectocarpus, she demonstrated that brown algae had two kinds of plurilocular organs: some occurring on hap- loid plants and functioning as gametangia and some on diploid plants and func- tioning as zoosporangia. The diploid plants frequently also formed unilocular sporangia. Meiosis occurred in the unilocular sporangia, as had previously also been shown by Yamanouchi with reference to Zanardinia and Cutleria and by Kylin (1918) with reference to Chorda. No reduction divisions occurred in the plurilocular sporangia of diploid plants and the zooids produced in them ger- minated directly. PAPENFUSS: CLASSIFICATION OF THE ALGAE 169 Knight (1923) showed that the life history of Pylaiella included an alterna- tion of isomorphic generations. She ( 1929 ) was unable, however, to demonstrate an alternation of generations in Ectocarpus. Contrary to the long-held view that the zooids from the unilocular organ were zoospores, she claimed that at least in British waters the zooids from the unilocular organs of Ectocarpus func- tioned as gametes. In this region there thus existed only diploid plants. She repeated the observations of Berthold and otliers at Naples and found that in that area the plants were haploid and their plurilocular organs were gamctangia. Papenfuss (1933, 1935), working at Woods Hole, Massachusetts, confirmed the observations of Knight that Ectocarpiis included haploid plants which bear only plurilocular organs and diploid plants which form both unilocular and plurilocular organs. He was unable, however, to confirm her observations re- garding the gametic nature of the zooids from the unilocular organs. Instead, he found that Ectocarpus exliibited a regular alternation of isomorphic genera- tions. The observations of Papenfuss were confirmed by F0yn (1934) working in Norway. Several other investigators have claimed a gametic role for the zooids from the unilocular organs of diverse brown algae. Although such behavior is theo- retically possible, the evidence presented for the alleged instances of conjuga- tion between these swarmers is not convincing. It would indeed be remarkable if an organism, such as Ectocarpus siliculosus, could form gametes on the diploid as well as the haploid generation. In 1904 AVilliams demonstrated the occurrence of an alternation of isomor- phic generations in Dictyota and in 1915 Sauvageau made the epoch-making discovery that Saccorhiza hulhosa, a member of the Laminariaceae, possesses an alternation of heteromorpliic generations comparable to that of ferns. The fa- miliar macroscopic plant was found to be the sporophyte. The zoospores formed in its unilocular sporangia give rise to microscopic, filamentous gametophytes which are dioecious and produce oogonia and antheridia. This very significant discovery of Sauvageau, which was made on the basis of cultures, created a great deal of interest in the brown algae. It was evident that the complete cycle of development of many of these algae could not be ascer- tained unless they were grown in culture. It was also clear that rich rewards were in store for those who would follow his approach to problems relating to the life histories of brown algae. He himself retained leadership in this fruitful field until his death in 1936. (For a list of his many publications see Dangeard, 1937.) The knowledge that has accumulated during the past fifty years has natu- rally had far-reaching effects on the classification of the Phaeophycophyta. Although Lamouroux (1813), C. Agardh (1817, 1824), and Harvey (1836) had recognized the autonomy of the brown algae, each of them had included in the group certain dark-colored red algae or had assigned representatives of the group to other major taxa. In 1848 J. Agardh published the first volume of his Species genera et ordines algarum, a volume devoted exclusively to the brown algae. The Phaeophyco- phyta were segregated by J. Agardh into seven tribes (also referred to by him in different places as families or orders), six of which were first recognized by Greville (1830) and Harvey (1836). It is noteworthy that each of these seven 170 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES groups was later elevated to the rank of order (with altered circumscription, of course). Between 1848 and 1917, these algae were divided by different authors into two, three, or four major taxa. Thuret (1855, pp. 5-15) recognized four groups: Phaeosporeae, Tilopterideae, Dictyoteae, and Fucaceae. Hauck (1883) recog- nized three orders: Fucoideae (with one family), Dictyotaceae (with one family), and Phaeozoosporeae (with ten families). Hauck 's three groups were retained by De Toni (1895) except that he used, respectively, the names of Cyclosporinae (a designation proposed by Areschoug, 1847, for the Fucaceae), Tetrasporinae, and Phaeozoosporinae. Kjellman (1891-1893) divided the Phaecophyceae into the two groups Phaeo- sporeae and Cyclosporeae (with the single family Fucaceae) and removed the Dictyotaceae to an independent group Dictyotales, which he considered as so different from other brown algae that they could not be properly placed with them (see also Falkenberg, 1882, pp. 169, 230-234). (Because they commonly form four immobile spores in their unilocular sporangia, which thus resemble the tetrasporangia of red algae, the Dictyotales were a stumbling block to many students of the algae of the last century.) The Phaeosporeae were divided by Kjellman into the two subgroups Zoogonicae and Acinetae (which included only the Tilopteridaceae). It is apparent that as far as the major categories are con- cerned Kjellman's system differed but little from that of Thuret. Oltmanns (1904) segregated the Phaeophyceae into the three groups Phaeo- sporeae, Akinetosporeae (a designation proposed by Bornet, 1891, p. 370), and Cyclosporeae. The Akinetosporeae received only the Tilopteridaceae (character- ized by their immobile monosporangia), but in contrast to earlier systems, Olt- manns placed the oogamous Dictyotaceae as a second family with the likewise oogamous Cyclosporeae (Fucaceae). In 1917 Kylin revised the classification of the Phaeophyceae, basing his sys- tem largely on developmental and nuclear cycles. He recognized five orders: Phaeosporeae, Tilopteridales, Dictyotales, Laminariales, and Fucales. The es- sentially new feature here is the establishment of the order Laminariales for those Phaeosporeae of earlier systems that had been found to possess an alter- nation of heteromorphic generations. During the next ten years the old order Phaeosporeae was further subdi- vided into the following seven orders: (1) Ectocarpales (Setcliell and Gardner, 1922, Oltmanns, 1922b; see Papenfuss, 1947, p. 398, fn., regarding the dates of the works by Setchell and Gardner, and Oltmanns); (2) Sphacelariales (Olt- manns, 1922b); (3) Cutleriales (Oltmanns, 1922b); (4) Chordariales (Setchell and Gardner, 1925); (5) Sporochnales (Sauvageau, 1926); (6) Desmarestiales (Setchell and Gardner, 1925); (7) Dictyosiphonales (Setchell and Gardner, 1925). Utilizing the recently acquired knowledge of the structure and reproduc- tion of the Phaeophycophyta, Kylin in 1933 erected a new system of classifica- tion of these algae. He divided them into three classes and a total of twelve orders, one of which (the Punctariales) he established in this paper as a segre- gate from the Ectocarpales. The first class, the Isogeneratae, received forms that showed an alternation of isomorphic generations. It included the orders Ecto- carpales, Sphacelariales, Cutleriales, Tilopteridales, and Dictyotales. The second PAPENFUSS: CLASSIFICATION OF THE ALGAE 171 class, the Heterogeneratae, comprised forms which showed an alternation of heteromorphic generations. It included two subclasses, the Ilaplostichineae and the Polystichineae. In the Ilaplostichineae no intercalary longitudinal di- visions occur in the thallus and consequently no true parenchymatous tissues are formed. In the Polystichineae intercalary longitudinal divisions are formed and hence true parenchymatous tissues are produced. The Ilaplostichineae re- ceived the orders Chordariales, Sporochnales, and Desmarestiales, whereas the Polystichineae received the Punctariales, Dictyosiphonales, and Laminariales. The third class, the Cyclosporeae, received the single order Fucales. The separation of the Ectocarpales sens^l Oltmanns (1922b) into haplosti- chous and polystichous groups was first proposed by Kuckuck {in Oltmanns, 1922b; 1929). It is to be noted, however, that Kylin employed this character only with reference to the Heterogeneratae. Papenfuss (1947) has merged the Punctariales in the Dictyosiphonales. It would seem that Kylin (1947) also arrived at the conclusion that these two orders are synonymous, since in the body of his paper he placed Bictyosiplwn in the Punctariales even though in the introduction (p. 4) he accepted both orders. Arasaki (1949) argues in favor of retention of both orders but the evidence produced is hardly sufficient. The system of Kylin has received wide recognition. Among those who have not accepted it or have accepted it only in part are Hygen, Fritseh, and more re- cently Papenfuss. Hygen (1934) is dissatisfied with Kylin 's arrangement largely because the Isogeneratae includes a heterogeneous assortment of algae that could not be regarded as forming a phylogenetically coherent unit. Fritseh (1943, 1944, 1945) does not accept the orders Chordariales, Punc- tariales, and Dictyosiphonales but retains the families comprising them in the Ectocarpales largely because he believes (1944, p. 254) that their heteromorphic life cycle is derived "by divergent development of the two generations, from an isomorphic alternation, comparable to that exhibited by the Ectocarpaceae." It seems very likely, however, that the other groups with a heteromorphic alterna- tion of generations (the oogamous Sporochnales, Desmarestiales, and Lamina- riales) also evolved, even if not directly, from the Ectocarpales. The Ectocar- pales sensu Fritseh includes a very heterogeneous assemblage of algae. Papenfuss (1951b) accepts all the orders recognized by Kylin, except the Punctariales, but rejects the classes Isogeneratae, Heterogeneratae, and Cyclo- sporeae and the subclasses Ilaplostichineae and Polystichineae of the Hetero- generatae. Such an arrangement allows for the parallel and independent evolu- tion of groups with an alternation of isomorphic or heteromorphic generations; it recognizes the Ectocarpales as the possible ancestral stock from which had emerged several orders; and it takes cognizance of the fact that the Fucales are parenchjnnatous (polystichous) and that parenchymatous forms also occur in the Isogeneratae (e.g., Dictyotales, Sphacelariales). In agreement with K.ylin (1933, 1937c, 1938, 1940b) and Fritseh (1945, pp. 380-381), Papenfuss regards the Fucales as occupying an isolated position in the Phaeophyceae. Recently Feldmann (1949) established an order Scytosiphonales for certain Dictyosiphonales, but the more significant distinguishing features of the new order are based on the acceptance of observations of extremely questionable accuracy. On the basis of pigment composition, the Phaeophycophyta appear to be 172 ^ CENTURY OF PROGRESS IN 7HE NATURAL SCIENCES related, even if only remotely, to both the Chrysophycophyta, especially the Ba- eilariophyceae, and the Pyrrophycophyta (cf. Strain, 1951, p. 253). The evi- dence derived from the structure of tlie flagella in these three groups (Petersen, 1918, 1929; Vlk, 1931, 1938; Deflandre, 1934; Longest, 1946; Koch, 1951; Man- ton and Clarke, 1951a, 1951b) suggests that a closer relationship exists between the Phaeophycophyta and Chrysophycophyta than between either of these two groups and the Pyrrophycophyta. The following arrangement of the Phaeophycophyta is essentially that of Papenfuss (1951a). Phylum PHAEOPHYCOPHYTA Papenfuss (1946, p. 218) Syn.: Phaeophyta Wettstein (1901, p. 46) Class Phaeophyceae De Bary (1881, p. 14) Syn.: Melanospermeae Harvey (1836, p. 157); Melanophyceae Rupreclit (1851, p. 206) ; Phycopheinophycees Marchand (1895, p. 15) Order ECTOCARPALES Setchell et Gardner (1922, p. 403) Family Ectocarpaceae (C. Agardli) Kiitzing orth. mut. Harvey (1849, p. 11) Syn.: Streblonemaceae Kylin (1947, p. 45), nomen nudum; Acineto- sporaceae Bornet orth. mut. Hamel (1931: p. 8; cf. Kornmann, 1953) Family Ralfsiaceae Hauck (1883, p. 318) Syn.: Lithodermataceae Hauck (1883, p. 318); Strangulariaceae Stromfelt (1886, p. 49); Nemodermataceae Feldmann (1937, p. 121) Order SPHACELARIALES Oltmanns (1922b, p. 83) Syn.: Discosporangiales 0. C. Schmidt (1937a, p. 3) Family Sphacelariaceae J. Agardh orth. mut. Cohn (1872a, p. 17) Family Stypocaulaceae Oltmanns (1922b, p. 95) Family Cladostephaceae Oltmanns (1922b, p. 102) Family Choristocarpaceae Kjellman (1S91, p. 190) Syn.: Discosporangiaceae 0. C. Schmidt (1937a, p. 3) Order CUTLERIALES Oltmanns (1922b, p. 109) Family Cutleriaceae (Thuret) Hauck (1883, p. 318) Order TILOPTERIDALES Kylin (1917, p. 308) Family Tilopteridaceae (Thuret) Cohn orth. mut. De Toni (1891b, p. 182) fFamily Masonophycaceae O. C. Schmidt (1937b, p. 5) Order DICTYOTALES Kjellman (1893, p. 291) Family Dictyotaceae Lamouroux orth. mut. Dumortier (1829, p. 76) Syn.: Zonariaceae (S. F. Gray) Nageli (1847, p. 179) Order CHORDARIALES Setchell et Gardner (1925, p. 570) Family Myrionemataceae (Nageli) Foslie orth. mut. Skottsberg (1907, p. 49) Family Elachistaceae Kjellman (1890, p. 41) Family Corynophlaeaceae Oltmanns (1922b, p. 23) Syn.: Leathesiaceae Setchell et Gardner (1925, p. 507) Family Chordariaceae (C. Agardh) Greville orth. mut. Harvey (1849, p. 11) Syn.: Mesogloeaceae Kiitzing (1843, p. 32); Myriogloiaceae Kuckuck ex Setchell et Gardner (1925, p. 555) ; Heterochordariaceae Setchell et Gardner (1925, p. 549) ; Aegiraceae Setchell et Gardner (1925, p. 543) ; Myriocladiaceae Kuckuck (1929, p. 63) Family Spermatochnaceae Kjellman (1890, p. 32) Syn.: Stilophoraceae (Nageli) De Toni et Levi orth. mut. Kjellman (1890, p. 34) Family Acrothrichaceae (Oltmanns) Kuckuck (1929, p. 66) Family Chordariopsidaceae Kylin (1940a, p. 53) Family Splachnidiaceae Mitchell et Whitting (1892, p. 9) PAPENFUSS: CLASSIFICATION OF THE ALGAE 173 Order SPOROCHNALES Sauvageau (1926, p. 364) Family Sporochnaceae Greville orth. mut. Harvey (1849, p. 10) Order DESMARESTIALES Setchell et Gardner (1925, p. 554) Syn.: Arthrocladiales Sauvageau (1931, p. 117) Family Arthrocladiaceae Chauvin orth. mut. Hauck (1883, p. 317) Family Desmarestiaceae (Thuret) Kjellman (1880, p. 10) Order DICTYOSIPHONALES Setchell et Gardner (1925, p. 586) Syn.: Punctariales Kylin (1933, p. 93) ; Scytosiphonales Feldmann (1949, p. 112) Family Striariaceae Kjellman (1890, p. 53) Syn.: Stictyosiphonaceae (Kjellman) Kuckuck (1929, p. 80) Family Giraudyaceae (Kjellman) Hygen (1934, p. 210) Family Myriotrichiaceae Kjellman (1890, p. 46) Family Punctariaceae (Thuret) Kjellman (1880, p. 9) Syn.: Encoeliaceae (Bory) Kiitzing orth. mut. Kjellman (1890, p. 55); Asperococcaceae (Zanardini) De Toni et Levi orth. mut. Foslie (1890, p. 88); Litosiphonaceae Kuckuck (1929, p. 80), nomen nudum; Soran- theraceae Kuckuck (1929, p. 80), nomen nudum Family Scytosiphonaceae (Thuret) Hauck (1883, p. 317) Syn.: Phaeosaccionaceae Feldmann (1949, p. 112) Family Chnoosporaceae Setchell et Gardner (1925, p. 552) Family Dictyosiphonaceae Kiitzing orth. mut. Kjellman (1890, p. 49) Syn.: Coilodesmaceae (Kjellman) Setchell et Gardner (1925, p. 577) Order LAMINARIALES Kylin (1917, p. 308) Family Chordaceae Dumortier (1822, pp. 72, 102) Family Laminariaceae (C. Agardh) Dumortier orth. mut. Dumortier (1829, p. 77) Syn.: Phyllariaceae (Kjellman) Hamel (1938, p. 304) Family Lessoniaceae (Setchell) Setchell et Gardner (1925, p. 621) Family Alariaceae (Kjellman) Setchell et Gardner (1925, p. 633) ^Family Prototaxitaceae Pia (1927, p. 95) Order FUCALES Kylin (1917, p. 309) Family Ascoseiraceae Skottsberg (1907, p. 148) Family Durvilleaceae (Oltmanns) De Toni (1891b, p. 173) Family Notheiaceae Kuckuck (1929, p. 12) ex 0. C. Schmidt (1938, p. 224) Syn.: Hormosiraceae (Gruber) Fritsch (1944, p. 257) Family Fucaceae Lamoroux orth. mut. Dumortier (1822, p. 72) Family Himanthaliaceae (Kjellman) De Toni (1891b, p. 173) Family Cystoseiraceae Kiitzing orth. mut. De Toni (1891b, p. 173) Family Sargassaceae (Decaisne) Kiitzing orth. mut. De Toni (1891b. p. 174) Phylum Schizopiiyta class schizophyceae Characterization: As a class the Schizophyceae (Cyanophyceae, Myxophyceae) or bluegreen algae as they are commonly called are sharply distinguished from the other large groups of algae. The low state of cell differentiation, the bluegreen color of the thalli, the production of cyanophycean starch (Kylin, 1943a) and the absence of an organized nucleus are characters that clearly set them apart. There is at present no good evidence indicating that they evolved from flagellated ancestors or that they are the direct ancestors of other algal groups, although a very distant relationship with the red algae is not improbable. The simplest Schizophyceae are unicellular. In many instances, however, the indi- vidual cells remain attached to one another to form colonies of various shapes and sizes. The more advanced types are filamentous. The filaments are simple or branched and may be aggregated. The branched forms exhibit false or true branching or both types of branching. The individual cells of the unicellular forms and the filaments of the fila- 174 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES mentous species are usually enveloped by a gelatinous sheath that may be homogeneous or stratified and frequently is pigmented. The inner portion of the sheath contains a small amount of cellulose (Kylin, 1943a). In the majority of bluegreen algae, two regions are distinguishable within the proto- plast: a peripheral region, the chromoplasm, which contains the pigments, and a colorless central region, the central body or centroplasm. Ordinarily the protoplast contains no evident vacuoles. Recent work indicates that the pigments in the chromoplasm occur in small bodies (Calvin and Lynch, 1952). The pigments consist as far as known of chlorophyll a, beta- carotene and another carotene (flavacin) found only in Schizophyceae, two xanthophylls that are peculiar to these algae, and two proteinaceous pigments (phycobilins), the blue c-phycocyanin and the red c-phycoerythrin (Strain, 1951, p. 253). The centroplasm contains various kinds of bodies, including some that are in the form of granules, rods, or reticula and become evident after application of the Feulgen nuclear reaction. The bodies are not bounded by a nuclear membrane, however, and no nucleoli appear to be present. In the Stigonemataceae and certain Scytonemataceae adjoining cells are united by pit connections. Only one such connection is present between any two cells and it is always a primary pit connection. All of the filamentous bluegreen algae, with the exception of the Oscillatoriaceae, regularly form a special type of cell known as a heterocyst. They originate from vege- tative cells and have a thickened wall. Intercalary heterocysts have a conspicuous pore at each end; terminal heterocysts have a pore only at the proximal end. The filaments frequently break at the heterocysts and these structures indirectly function in vegetative multiplication. In some instances they have been observed to produce new filaments. (See Fritsch, 1951, for a discussion on these cells.) As far as known sexual reproduction does not occur in the Schizophyceae. Vegetative multiplication by fission in the unicellular or colonial forms is of common occurrence. In the order Hormogonales the chief method of multiplication is by short lengths of the vegetative filaments called hormogonia. The hormogonia are delimited by the death of occasional cells at intervals along the length of the filament. In certain forms the hormo- gonia are modified as organs of perrenation (hormocysts). Many of the filamentous genera, but no Oscillatoriaceae, form thick-walled resting spores known as akinetes. In certain genera of the Chamaesiphonales the contents of a cell divide into a number of endospores. These spores are thin-walled and germinate directly to produce new plants. The bluegreen algae live in many kinds of habitats. Many are aquatic in freshwater or marine situations; others are terrestrial or subaerial in occurrence. A number of forms grow in hot springs, at times with a temperature as high as 85° C. Many forms live in association with other organisms — plants and animals. Species belonging principally to the genera Gloeocapsa, Nostoc, Scytonema, and Stigonevia constitute the algal associate in many lichens. A number of species are able to fix atmospheric nitrogen. History: Although members of this class have been known to the world of science since the time of Linnaeus (1753), who described a few species under the generic names Tremella, Byssus, and Viva, the distinctive features of the class remained unrecognized until the middle of the nineteenth century. To be true C. Agardh in 1824 established an order Nostochinae (one of six into which he divided the algae) to receive Nostoc and Bivularia, but he also assigned to this order several genera belonging to unrelated groups of algae and referred other genera of bluegreen algae (e.g., OsciUatoria) to the order Confervoideae. The bluegreen algae were first recognized as constituting an autonomous group (order) of algae by Stizenberger in 1860 (p. 18). He called them Myxo- phyceae, adopting a designation (Myxophykea) previously used by Wallroth (1833, p. ix) for a heterogeneous assemblage of algae, including representatives PAPENFUSS: CLASSIFICATION OF THE ALGAE 175 of the Scliizophyceae. Stizenberger remarked that these algae were distinguish- able from other algal orders by their pigments. Rabenhorst (1863, p. 1) established a division for these algae which, on the basis of color, he named Phj^cochromaceae. As one of the features of the group, Rabenhorst mentioned the absence of a nucleus in the cells, a character pre- viously remarked upon by Nageli (1849, p. 45) with reference to some of the unicellular members of the assemblage. Rabenhorst credited the group with six families, viz., Chroococcaceae, Os- cilla[to]riaceae, Nostocaceae, Rivulariaceae, Scytonema[ta]ceae, and Sirosi- phonaceae (= Stigonemataceae), all of which are still accepted, although usually with modified circumscriptions. In 1865 Rabenhorst regarded tlie bluegreen algae as a class and changed the divisional name Phycochromaceae, which he had given them in 1863, to Phy- cochromophyceae. At this time Rabenhorst divided the class into two orders: "Ordo I. Cystiphorae," in which he placed the family Chroococcaceae, comprised of unicellular and colonial forms, and "Ordo II. Nematogenae," to which he referred the remaining five families of his treatment of 1863, all of which in- cluded filamentous forms. On account of their blue pigment, Sachs (1874) named these algae Cyano- phyceae. Because of the appropriateness of this designation and because of the fame of Sachs and his textbook, the name Cyanophyceae immediately gained favor among botanists, and it is still used by many. Cohn (1880, p. 286) gave these algae the name Schizophyceae.^*' He re- garded them as a group coordinate with the bacteria (which he in 1872 [a] had named Schizomycetae) and placed both groups in his order Schizosporeae, as he had first done in 1872 (a and b). That there existed a relationship between these two groups of organisms had already been pointed out by Cohn as early as 1854. Engler (1892) divided the thallophytes into two divisions, the Myxothallo- phyta and the Euthallophyta. Under the latter he had a number of subdivisions, one of which was Schizophyta, to which he referred the two classes Schizophy- ceae and Schizomycetes. Without giving the reference, Pringsheim (1949, p. 48) and others credit Cohn with the name Schizophyta, but I have been un- able to find this designation in Cohn's publications. It apparently was used for the first time by Falkenberg (1882, p. 162) as the name of an order com- prising both the bluegreen algae and the bacteria. In conformity with the views of Cohn and the system of Engler, which has had many adherents down to the present, the bluegreen algae are here regarded as constituting a class, the Schizophyceae, coordinate with the bacteria (class Schizomyceteae) in the phylum Schizophyta. It should be pointed out, however, that many biologists do not believe that these two (or several) groups of organ- isms are related. This latter view has been particularly well defended by Prings- heim (1949), to whom the reader is referred for a detailed discussion of the question (see also Stanier and van Niel, 1941). Although the characters which point to an af^nity between bacteria and 10. It is to be noted, however, that Rabenhorst (1847, pp. V, 16) had previously used the designation Schizophyceae for a "suborder" of algae comprising the diatoms and desmids. 176 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES bluegreen algae are largely of a negative kind, as Pringsheim has emphasized, there is little reason for believing that certain negative characters are less im- portant than positive characters as indicators of phylogenetic relationship. Fur- thermore, if present-day bacteria and bluegreen algae had evolved from com- mon ancestors affinities should be sought primarily among the simpler members of both groups (the Eubacteriales and the Chroococcales, respectively), since they are the forms which may be expected to have retained and consequently show the largest number of ancestral characters. The morphological similarity between certain members of the Eubacteriales and members of the Chroococ- cales suggests that a relationship does exist between these two groups. Further evolution in the bluegreen algae has resulted in the development of thalli which are much more elaborate than those of the higher bacteria. But morphological differences of comparable magnitude are not uncommon among groups of organ- isms which are known to be phylogenetically related and their occurrence in the Schizophyta do not necessarily speak against a common origin of bac- teria and bluegreen algae. On the basis of method of multiplication, Thuret as long ago as 1875 subdi- vided the bluegreen algae, or Nostochinees as he called them, into two tribes: (1) the Chroococcaceae or Coccogoneae, which show vegetative propagation by single cells, and (2) the Nostochineae or Hormogoneae, which reproduce vege- tatively by short rows of cells (hormogonia). Thuret segregated the Hormo- goneae into two subtribes: (1) Psilonemeae, in which the filaments lack hairlike tips, and (2) Trichophoreae, in which the filaments possess hairlike apices. In Thuret 's time the members of the subsequently established family Chamaesi- phonaceae were only poorly known. To Nageli (1849) and Hansgirg (1888b, 1892) we are indebted for much of the fundamental information on which present-day classification of the Chroo- coccaceae rests. Current classification of Thuret 's Hormogoneae is largely based on the systems of Borzi (1878, 1879, 1882), Bornet and Flahault (1886-1888), and Gomont (1892, 1893). The first comprehensive treatment of the Schizophyceae as a whole was given by Kirchner (1898), who followed, as far as the broad outlines are concerned, the classification of Rabenhorst (1865), Thuret (1875), and Hansgirg (1888b, 1892). In accordance with the system of Thuret, Kirchner divided the blue- green algae into Coccogoneae and Hormogoneae. In the Coccogoneae he placed the Chroococcaceae and the Chamaesiphonaceae, a family established by Borzi in 1882. The Hormogoneae were segregated, in agreement with Thuret, into the Psilonemateae (which received the families, Oscillatoriaceae, Nostocaceae, Scytonemataceae, and Stigonemataceae) and the Trichophoreae (in which were placed the Rivulariaceae and the Camptotrichaceae). As has been pointed out by Fritsch (1944, p. 262), this division of the Hormogoneae (^ Hormogo- nales) by Thuret and Kirchner into two groups on the absence or presence of hairs at the tips of the filaments overemphasized the systematic value of a minor character, and is no longer adhered to. Adopting and amending a classification introduced by Stizenberger (1860), Bornet and Flahault (1886, p. 325) and Gomont (1892) divided the Hormo- goneae into the two groups Heterocysteae and Homoej^steae according as the trichomes contain or lack heterocysts. Kirchner (1898, p. 49) and Fritsch (1944) PAPENFUSS: CLASSIFICATION OF THE ALGAE 177 have emphasized, however, that rehited genera are segi-egated l)y tliis division. In more recent times the division of the Hormogonales into Ileterocysteae and Homocysteae has been followed by Setchell and Gardner (1919) and by Smith (1933) who, however, has since abandoned this classification (Smith, 1950). In 1895 ]\Iarchand established the ordinal names Coccogonees and Hormo- gonees (changed to Coceogonales and Hormogonales by Atkinson, 1905, p. 163) for the two tribes of Thuret. The name Hormogonales is still accepted by many phycologists but the designation Coceogonales has been abandoned in favor of Chroococcales, which was proposed by Wettstein (1924). Borzi had already in 1878 divided the order Nematogenae of Rabenhorst into the two suborders Hormogoneae and Cystogoneae. The latter suborder included only his new family Chamaesiphonaceae (p. 238) whereas the former received the Nostocaceae, Scytonemataceae, Rivulariaceae, and Oscillatoriaceae. Borzi placed, as others before him had done, the Chroococcaeeae in an order by itself, which he called Gloeogenae. Following up the train of thought of Borzi, Hans- girg (1892, p. 17) divided the bluegreen algae into the three orders Gloeosipheae (= Hormogonales), Chamaesiphonaceae, and Chroococcoideae. The order Cha- maesiphonaceae received attached, unicellular (Chamaesiphon) or filamentous {Clasiidium) forms, which occur as solitary individuals or as colonies, lack hor- mogonia and heterocysts, and multiply by spores (endospores) produced in ba- sipital succession. Finally in 1924 Wettstein proposed the currently accepted designation Cha- maesiphonales for the family Chamaesiphonaceae. In his treatment of the Schizophyceae in De Toni's Sijlloge algarum, Forti (1907) followed the classification of Kirchner. Borzi, shortly afterward in a series of papers (1914, 1916, 1917), presented a revision of his earlier (1878, 1879, 1882) classification of these algae. Some of the new features of this sys- tem were later adopted by Geitler in the development of his system. By the year 1925, the Schizophyceae had thus by degrees come to be segregated into three orders and a total of 14 families, including one (Microchaetaceae) w^hich had been erected by Lemmermann in 1907, two (Hyellaceae and Borziaceae) which were established by Borzi in 1914, and three (Nodulariaceae, Lepto- basaceae and Loefgreniaceae) which were created by Elenkin in 1916 and 1917. In 1925 (a and b) Geitler published a system which formed a radical depar- ture from previous classifications. He divided the bluegreen algae into seven orders and a total of 19 families. Shortly afterward Geitler (1930-1932) aban- doned in part his sj'stem of 1925 and recognized only the three old orders Chroo- coccales, Chamaesiphonales, and Hormogonales. At this time Geitler regarded the Schizophyceae as comprised of 21 families, the majority of which were the same as those accepted in 1925 but some of the ones recognized in 1925 were reduced and several new ones were added. In 1942 (pp. 37 ff.) Geitler returned in part to his system of 1925 and recognized four orders (Chroococcales, Pleuro- capsales, Dermocarpales, Hormogonales) and 22 families. In 1938 and 1949 appeared the first and second fascicles, respectively, of the systematic part of Elenkin 's monumental work on the freshwater and terrestrial Schizophyceae of Russia. Elenkin elaborated upon Geitler's systems of 1925 and 1930-1932, and recognized, as far as the groups under consideration by him- self were concerned, no fewer than 12 orders and 47 families. 178 ^ CENTURY Of PROGRESS IN THE NATURAL SCIENCES It is to be regretted that both Geitler and Elenkin have burdened the already involved nomenclature of the Schizophyceae with a number of unnecessary names. These authors have violated the Code by renaming families whose cir- cumscription they have changed but which still include the type of the rejected family or, in the case of Geitler, by renaming families if the generic name from which a family name was derived has been reduced to synonymy. Fritsch (1942, 1944, 1945) accepts in part Geitler's system of 1925 and di- vides the Schizophyceae into the five orders Chroococcales, Chamaesiphonales, Pleurocapsales, Nostocales and Stigonematales. In the division of the class into five orders, the system of Fritsch corresponds closely to that of Geitler as amended in 1942, except that Geitler at this time maintained the order Hormo- gonales as a single taxon (as he had also done in 1930-1932) whereas Fritsch recognizes in its place the two orders Nostocales and Stigonematales, as Geitler had in 1925. Fritsch (1945) arranges the genera in 19 families, all of which, with the ex- ception of the Cyanochloridaceae and Loefgreniaceae, were recognized also by Geitler (1942), although the two authors do not always use the same names or place the families in the same order. Fremy (1930, 1933), Copeland (1936), Huber-Pestalozzi (1938), Lindstedt (1943), Skuja (1948), Smith (1950), Prescott (1951), and others accept only the three original orders Chroococcales, Chamaesiphonales, and Hormogonales, except that Copeland and Smith use the name Oscillatoriales Copeland (1936) instead of Hormogonales. Drouet (1951), however, recognizes no orders in the bluegreen algae and accepts only eight families. Evidently little agreement exists among students of the Schizophyceae as regards the classification of the class. This disagreement is attributable not so much to lack of knowledge of the morphology of these algae (although it seems likely that cultural studies will yield information that will be useful in the taxonomy of the group) as it is to the paucity of sharply defined characters and the existence of intermediate types which preclude the establishment of clear- cut taxa. The wide divergence in the systems proposed by the various specialists on the group hinges primarily on the taxonomic value assigned to the available characters. In his recognition of only eight families and the suppression of all orders, Drouet is probably guided by the existence of transitional types, al- though he has not yet presented the detailed arguments upon which his deci- sions are based. The separation of the class into the three orders Chroococcales, Chamaesi- phonales, and Hormogonales takes account of the structure of the thalli (uni- cellular or colonial or pseudofilamentous in the Chroococcales, unicellular or pseudofilamentous or filamentous in the Chamaesiphonales, multicellular and filamentous in the Hormogonales) and the method of multiplication (vegeta- tively by cell division or colony fragmentation in the Chroococcales, by endo- spores in the Chamaesiphonales, by hormogonia and in some instances by aki- netes in the Hormogonales). The division of the Chamaesiphonales into the two orders Chamaesiphonales and Pleurocapsales by Fritsch (1942, 1944, 1945) and Geitler (1942, as Dermo- earpales and Pleurocapsales) takes cognizance of differences in thallus organi- zation— the Chamaesiphonales receiving plants which are unicellular and with PAPENFUSS: CLASSIFICATION OF THE ALGAE 179 the cells exhibiting polarity as contrasted with the filamentous and frequently heterotrichous habit of the forms placed in the Pleurocapsales. It is to be noted, however, that these two authors do not in all instances agree in their assign- ment of families to these two orders. The separation of the Hormogonales into Nostbcales and Stigonematales by Fritsch is based on the occurrence of true branching and the heterotrichous habit of the thalli in some forms (Stigonematales) as contrasted with the un- branched or falsely branched condition of the filaments in others (Nostocales). The division of the bluegreen algae by Elenkin into a large number of orders and families is an attempt to segregate the genera on the basis of small differ- ences into seemingly clear-cut groups. Elenkin (1933) thus, for example, ele- vates the Chroococcaceae to the rank of order and divides it into ten families on the basis of the planes of division of the cells and the geometric form of the colonies. Subdivision to the extent proposed by Elenkin is probably unwar- ranted since it removes from one another forms which seemingly are so closely related that some authors (e.g., Drouet and Daily, 1952) reduce them to a bare few genera and species. The following synoptic arrangement of the orders and families is the author's compromise of the various recent systems of classification of the Schizophyceae^^. According to the current Code, the nomenclature of the Chroococcales and Cha- maesiphonales starts with Linnaeus (1753), that of the Oscillatoriaceae (Hor- mogonales) with Gomont (1892-1893), and that of all other Hormogonales with Bornet and Flahault (1886-1888). Phylum SCHIZOPHYTA (Falkenberg) Engler (1892, p. 3) Class ScHizopHYCEAE Colm (1880, p. 286) Syn.: Division Phycochromaceae Rabenhorst (1863, p. 1); Class Phycochromo- phyceae Rabenhorst (1865, p. 1): Order Myxophyceae Stizenberger (1860, p. 18); Cyanophyceae Sachs (1874, pp. 248, 251); Phycocyanophycees Marchand (1895, p. 11.) Non Schizophyceae Rabenhorst (1847, pp. v, 16) Order CHROOCOCCALES Wettstein (1924, p. 79) Syn.: Entophysalidales Geitler (1925a, p. 223); Tubiellales Elenkin (1934, p. 56); Coccogonales (Thuret) Marchand orth. mut. Atkinson (1905, p. 163) Family Chroococcaceae Nageli (1849, p. 40) Syn.: Coccobactreaceae Elenkin (1933, p. 19); Beckiaceae Elenkin (1933, p. 19); Merismopediaceae Elenkin (1933, p. 19); Microcysti- daceae Elenkin (1933, p. 19); Gloeocapsaceae Elenkin (1933, p. 19); Coelosphaeriaceae Elenkin (1933, p. 19) ; Gomphosphaeriaceae Elenkin (1933, p. 19) ; Woronichiniaceae Elenkin (1933, p. 19); Holopediaceae Elenkin (1933, p. 19) ; Cyanidiaceae Geitler (1933, p. 624) Family Entophysalidaceae Geitler (1925a, p. 235) Syn.: Chlorogloeaceae Geitler (1925a, p. 235) ; Tubiellaceae Elenkin (1934, p. 56) Order CHAMAESIPHONALES Wettstein (1924, p. 79) Syn.: Pleurocapsales Geitler (1925a, p. 238); Dermocarpales Geitler (1925a, p. 238); Siphononematales Geitler (1925a, p. 238); Endonematales Elenkin (1934, p. 57) Family Pleurocapsaceae Geitler (1925a, p. 238) Syn.: Chroococcidiaceae Geitler (1933, p. 623); Xenococcaceae Erce- govic (1932, p. 138); Podocapsaceae Ercegovic (1932, p. 138) 11. Several groups of organisms (other than the Beggiatoaceae) which have been placed with the bacteria (Achromatiaceae, Vitreoscillaceae, Thriotrichaceae, Cyanochlo- ridaceae) but which may be bluegreen algae, or at least include forms which probably are bluegreen algae, are discussed by Pringsheim (1949). 180 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Family Hyellaceae Borzi (1914, p. 359) Syn.: Scopulonemataceae Ereegovic (1932, p. 138) Family Dermocarpaceae Geitler (1925a, p. 247) Family Clastidiaceae Drouet et Daily (1952, p. 223) Family Chamaesiphonaceae Borzi (1882, p. 298) Family Siphononemataceae Geitler (1925a, p. 251) Family Endonemataceae Pascher (1929b, p. 347) Syn.: Pascherinemataceae Geitler (1942, p. 99) Order HORMOGONALES (Thuret) Marchand orth. mut. Atkinson (1905, p. 163) Syn.: Gloeosiphonales Wettstein (1924, p. 80); Oscillatoriales Copeland (1936, p. 78); Nostocales Geitler (1925a, p. 252); Stigonematales Geitler (1925a, p. 252); Mastigocladales Elenkin (1938, p. 528); Diplonematales Elenkin (1938, p. 535) Family Oscillatoriaceae (S. F. Gray) Dumortier ex Kirchner (1898, p. 61) Syn.: Borziaceae Borzi (1914, p. 358); Beggiatoaceae (Hansg.) Migula (1895, p. 41) ; Pseudonostocaceae Elenkin (1949, p. 1222) ; Schizo- thrichaceae Elenkin (1949, p. 1668) ; Crinaliaceae Elenkin (1949, p. 1845); Vaginariacees (Gomont) Marchand (1895, p. 12); Lyngbyacees Kiitzing orth. mut. Marchand (1895, p. 12) Whether the Beggiatoaceae are colorless bluegreen algae or bacteria has been a matter of disagreement for a long time. The evidence in favor of placing them in the Schizophyceae is presented by Pringsheim (1949), Family Gomontiellaceae Elenkin (1936, p. 543) Family Nostocaceae Dumortier ex Engler (1892, p. 4) Syn.: Anabaenaceae Elenkin (1938, p. 643); Aphanizomenonaceae Elenkin (1938, p. 845) Family Microchaetaceae Lemmermann (1907, p. 101) Syn.: Nodulariaceae Elenkin (1916, not seen, cited from Geitler, 1942, p. 159) ; Leptobasaceae Elenkin (1917, p. 164) Family Rivulariaceae Kiitzing ex Bornet et Flahault (1886, p. 338) Syn.: Camptotrichaceae (West et West) Kirchner (1898, p. 90); Til- deniaceae Kossinskaja (1926, p. 82); Hammatoideaceae Elenkin (1949, p. 1806); Homoeothrichaceae Elenkin (1949, p. 1813);? Sokoloviaceae Elenkin (1926, pp. 93, 95; 1949, p. 1834; cf. Geitler, 1942, p. 176) Family Scytonemataceae Kiitzing ex Bornet et Flahault (1887, p. 81) Syn.: Hydrocoryuaceae Elenkin (1949, p. 991); Plectonemataceae Elenkin (1949, p. 1772) ; Pseudoscytonemataceae Elenkin (1949, p. 1805); Pseudodiplonemataceae Elenkin (1949, p. 1838) Family Mastigocladaceae Geitler (1925a, p. 263) Syn.: Lithonemataceae Elenkin (1949, p. 185) Family Diplonemataceae (Borzi) Elenkin (1934, p. 79) Syn.: Borzinemataceae Geitler (1942, p. 141) Family Pulvinulariaceae Geitler (1925a, p. 254) Syn.: Loriellaceae Geitler (1925a, p. 253) Family Capsosiraceae Geitler (1925a, p. 255) Syn.: Pseudocapsosiraceae Elenkin (1949, p. 1849) Family Nostochopsidaceae Geitler (1925a, p. 257) Syn.: ?Loefgreniaceae Elenkin (1917, p. 161; cf. Geitler, 1942, p. 135) Family Mastigocladopsidaceae Iyengar et Desikachary (1946, p. 58) Family Stigonemataceae Kirchner (1898, p. 80) Syn.: Sirosiphonaceae (Stizenberger) Rabenhorst ex Engler (1892, p. 4) Phylum Rhodophycophyta Characterisation: The algae belonging to this phylum owe their characteristic red color to the presence in their plastids of a water-soluble proteinaceous pigment (phycobi- PAPENFUSS: CLASSIFICATION OF THE ALGAE 181 lin), r- phycoerythrin. Some forms contain, in addition, a second water-soluble proteina- ceous pigment, the blue r- phycocyanin. These two pigments commonly obscure the other pigments, which are: chlorophyll a, chlorophyll d, the xanthophyll lutein, alpha- and beta- carotene. A number of genera are colorless or nearly so and live as parasites on other red algae. In the simplest red algae the thallus consists of a single cell (Porphyridium, Chroo- theca). At the other extreme there are forms with a comparatively large, although never massive, foliaceous thallus (Iridaea, Aeodes). Flagellated vegetative or reproductive cells are entirely lacking in this group. Cells of red algae have a wall that is differentiated into an inner cellulosic and an outer pectic portion. Calcification of the wall occurs in the coralline algae and in a number of other genera belonging to the orders Cryptonemiales and Nemalionales. (Encrusting coralline algae assist immensely in the building of coral reefs and often play a more important part in this process than the corals themselves. Fossil corallines are known from the Cretaceous onwards.) In primitive forms the cells are uninucleate, in others they are uni- or multinucleate, although the more highly evolved forms are always multinucleate. The reproductive organs are almost always uninucleate. In the less specialized forms the cells usually contain a single or only a few plastids. In many of these forms the plastid is axile in position and more or less stellate in form. In the higher forms each cell usually contains several to many discoid, lenticular, or bandlike chromatophores. In the lower forms the plastids frequently contain pyrenoids which usually lack a starch sheath. The product of photosynthesis is a polysaccharide known as floridean starch. Growth of the thallus is diffuse in the Bangiophycidae, apical or marginal in the Florideophycidae. In the latter, adjacent cells are joined by pit connections. Sexual reproduction in the red algae is always oogamous. The female sex organ, known as the carpogonium, is usually borne at the end of a special filament, the car- pogonial branch, and it usually forms a receptive process, the trichogyne. Only one egg is formed in a carpogonium. It never retracts from the carpogonial wall to become an individualized egg, either before or after fertilization. The male sex organ, or sperma- tangium, forms a single motionless spermatium which is conveyed passively to the trichogyne. Following its fusion with the latter, the spermatial nucleus migrates down into the carpogonium where it fuses with the egg nucleus. In the Bangiophycidae the fertilized carpogonium by division gives rise directly to a number of carposporangia. In the Nemalionales it produces gonimoblast filaments (the cystocarp) which form carposporangia. In the majority of Florideophycidae above the Gelidiales a diploid nucleus is conveyed to one or more generative auxiliary cells from which the gonimoblast is produced. In the higher groups the carpospores produce free-living tetrasporangium- forming diploid plants that resemble the sexual plants. History: As was mentioned in the introduction to this chapter, Lamouroiix (1813) was the first to remove, on the basis of color, certain red algae (11 genera) from comparable morphological types of a different color. He created a special category ("ordre") for these plants and named it Floridees. Thus La- mouroux in effect became the founder of the phylum Rhodophycophyta, although the group did not receive this status until almost a century later. The Florideae, or Florideophycidae as they should be known in conformity with the current botanical code of nomenclature, still constitute one of the two subclasses of the class Rhodophyceae. Adopting the designation of Lamouroux, C. Agardh in 1817 made the Flori- deophycidae one of the five sections into which he divided the algae. Whereas Lamouroux and C. Agardh failed to distinguish sharply between green, brown, and red algae, Harvey's (1836) treatment of them in Mackay's Flora hihernica represents a more complete separation between these three major groups of algae. In only a few instances did he assign genera to the wrong color group. 182 ^ CENTURY Of PROGRESS IN THE NATURAL SCIENCES Harvey (1836) proposed the name Rhodospermeae for the red algae, which desig- nation was changed to Rhodophyceae by Rnprecht (1851, 1855). Kiitzing (1843, pp. 20, 21) suggested the names phycoerythrin and phycocyanin for the two phycobilin pigments present in the plastids of these algae. (For summaries of our knowledge of the pigments of red algae reference should be made to the reviews by Kylin, 1937a, and Strain, 1951). Since present-day classification of the red algae is to a large degree based on the details of development of the reproductive organs, emphasis will be placed in this brief review on the growth of our knowledge of the reproductive processes of the group. Ellis (1767) and C. Agardh (1828, pp. 57-58) referred to the clusters of spermatangia as male reproductive organs. C. Agardh used the term antheridia for those of Polysiphonia merely because of their superficial resemblance to an anther. That they indeed were male structures was first established by Bornet and Thuret (1866a, 1866b, 1867). That the same species of red alga may include two kinds of plants, each with its own kind of spore-bearing structure (cystocarp and tetrasporangium) was first emphasized by Stackhouse (1801, p. xxvi). At first. Turner (1802, pp. 293- 294) and others strongly opposed this view, believing that different species were involved, but later Turner (1808, p. 130) remarked about this phenomenon as follows: Of the zeal, with which the study of Marine Botany has been cultivated during the few years that have elapsed subsequently to the publication of the Nereis Britannica [by Stackhouse, 1795-1801], and the Synojysis of the British Fuci [by Turner, 1802], some idea may be formed from the circumstance of the double fruit of F. [ucus] coccineus l^Plocaviiiim vorciiieutn}, being at that time regarded as a curiosity, and as so extra- ordinary to be in itself almost sufficient to justify the dividing of the plant into two distinct species," whereas a similar appearance is now known to be observable in several of its congeners, and we have every reason to believe, that in the course of time it will be discovered in many others. In 1847 Harvey remarked (p. 4) : "The Ehodosperms are remarkable for possessing what seems to be a double system of fructification, a thing without parallel in the Vegetable Kingdom." On account of this feature, Kiitzing (1843) had previously named them Heterocarpeae. Decaisne (1842a) considered the tetrasporangium as the "typical" reproduc- tive organ of red algae and the cystocarp as a sort of proliferation or gemma. Harvey (1849, pp. 67-74), on the contrary, was of the opinion (p. 73) that the spores formed in the cystocarp should be considered ". . . of the nature of seeds [that is, the result of a sexual process], and not as huds," and that the spores formed in the tetrasporangium "should be regarded as gemmules." Be- cause the clusters of spermatangia occur in a position similar to that of the cystocarps in many genera of red algae (on trichoblasts in the Rhodomelaceae), Harvey argued that these structures (the "antheridia" of C. Agardh) might be of the nature of stamens. In the same work he remarked, however (p. 73) : ... we do not yet know the cause of the formation of conceptacles [cystocarps] and the production of spores. We know that seeds result from the joint agency of stamens and 12. Turner (1802) had on this account divided it into two varieties and remarked (p. 294), "There can indeed be but little question of their being in reality separate species . . ." PAPENFUSS: CLASSIFICATION OF THE ALGAE 183 pistils. But we do not know wliether any process similar to fertilization takes place with the spores of these algae. Niigeli (1847) divided the algae into two classes: (1) "Algae," which lack sexual reproduction, and (2) "Floridcae," which reproduce sexually. He re- garded the tetrasporangia of the Florideae as female sex organs which produce four spores, the antheridia of C. Agardh and others he considered male sex or- gans wliich produce sperms, and the cystocarps he regarded, in agreement with Decaisne, as structures of vegetative reproduction. Ruprecht (1851, pp. 205- 206), on the other hand, thought the tetraspores corresponded to the pollen and the carpospores ("Samen") to the seeds of phanerogams. He thought the an- theridia produced sperm cells, which were lacking in phanerogams, although this had not yet been established. It is evident that Ruprecht, like earlier botan- ists and those of the following fifty years, did not understand tlic role of the tetrasporangia in the life history of these algae. Thuret (1851) illustrated and described in unqualified terms as antheridia, structures which he studied in several red algae and their contents as anthero- zooids (a term proposed by Derbes and Solier, 1850, p. 263), although he was unable actually to determine their role inasmuch as he found that both the car- pospores and the tetraspores would germinate without having been in contact with the "antherozooids." Thuret 's observations were confirmed by Pringsheim (1855). Finally, Bornet and Thuret in 1866 and 1867 for the first time clearly de- scribed sexual reproduction in a number of red algae. They determined the na- ture of the female apparatus, which Nageli (1861) had observed but had misin- terpreted, and saw the spermatia attached to and coalescing with the trichogyne. Bornet and Thuret's discovery that the female gamete is produced in the ter- minal cell of a special filament, the carpogonial branch, and that tliis gamete is not liberated from the gametangium explained in large part why sexuality had eluded the various earlier investigators. From what was known about sexual reproduction in other groups, it was thought the female gamete would be an individualized protoplast like the egg of Fucus or of Volvox. The observations by Bornet and Thuret were extended by themselves (1876, 1878, 1880), Solms-Laubach (1867), Janczewski (1876), Schmitz (1879b, 1883) and others. Schmitz (1883) w^as the first to observe that in certain red algae the fertilized carpogonium produces filaments that fuse with a neighboring cell, which he (p. 229) termed the auxiliary cell, and that the gonimoblast develops from this cell. He thought that a second fusion of nuclei occurred in the auxil- iary- cell and that red algae consequently showed a double fertilization. He anticipated the skepticism that his interpretation of this phenomenon would generate, for he wrote (p. 246) : Einen zweimaligen Befruchtungsact im Entwickelungskreis einer einzelnen Species anziinehmen, dagegen straubt sich jedoch zur Zelt die botanische Anschauung voUstandig, das widerspricht aller Tradition. Oltmanns (1898) later showed that no fusion of nuclei occurs in the auxiliary cell, which receives a fusion (diploid) nucleus from the connecting filament but whose own (haploid) nucleus migrates to one side of the cell and plays no part in the ensuing development. 184 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES "VVille in 1894, working on Nemalion, saw the actual fusion of the male and female nuclei, an observation that was confirmed a few years later by Osterhout (1900). Yamanouehi (1906a, 1906b) first worked out the nuclear cycle and showed that the red algal genus Polysiphonia possesses an alternation of genera- tions between haploid gametophytic and diploid tetrasporangial plants, with meiosis occurring in the young tetrasporangium. Thus at last was determined the long-misunderstood role of the tetrasporangia in the life history of these plants. The correctness of Yamanouchi's observations was confirmed by Lewis (1912) by cultural studies. Svedelius (1915), studying the development and cytology of Scinaia, a genus which was known to lack tetrasporangia, established that in it meiosis occurs immediatel}^ after fertilization. Such species (the majority of Ne- malionales) consequently lack free-living tetrasporophytes. The observations of Svedelius were quickly confirmed by Kylin (1916) and Cleland (1919). A number of Florideophycidae — members of both the Nemalionales and of some of the orders above them — -were later found to have life cycles that deviate from the two general types referred to above. For a review of these the reader is re- ferred to the papers by Drew (1944) and Papenfuss (1950b). Oltmanns (1898, p. 138; 1904, p. 537) in agreement with Harvey (1849) and certain other early writers regarded the tetrasporangia as accessory repro- ductive organs. He considered the plant that produces the sex -organs as the gametophyte and the gonimoblast as the sporophyte (the carposporophyte of Church, 1919b, p. 331). From the cytological work of Yamanouehi, Lewis (1909), and many later investigators, it is now well established that the ma- jority of red algae above the Nemalionales possess three generations: a haploid gametophyte, a diploid carposporophyte which is permanently attached to and largely parasitic on the gametophyte, and a diploid, free-living tetrasporophyte. For an interesting account of the history of the discovery of an alternation of generations in the red algae the reader is referred to a paper by Svedelius (1916). Feldmann (1952) is of the opinion that all Florideophycidae that lack a free- living tetrasporophyte are derived. Although this is unquestionablj^ true of a number of forms — for example, certain species of Phyllopliora, Gymnogongrus, and Ahnfeldtia — it may be questioned whether this is also true of those Nema- lionales in which meiosis occurs immediately after fertilization (the majority of species in the order) or at carpospore formation. Svedelius (1953, p. 80, fn.) has promised to deal with this question. Although credit must go to Lamouroux (1813) for first recognizing the red algae as an autonomous group of plants, he, like his predecessors (e.g., Gmelin, 1768), and contemporaries (e.g., Esper, 1797-1808, and Turner, 1802, 1808, 1809, 1811, 1819), did not depart from Linnaeus (1753) in classifying these algae almost entirely on their external features, even if in much greater de- tail than Linnaeus. This very frequently resulted in the placing together of totally unrelated forms or in the separation of related forms. C. Agardh (1824, 1828) was the first to take into serious consideration in the classification of the algae the structure of the thallus and of the reproduc- tive organs, even if only as regards their gross structure. With C. Agardh thus begins what Sjostedt (1926, p. 78) has termed the anatomical period in the PAPENFUSS: CLASSIFICATION OF THE ALGAE 185 classification of the red algae and other groups. In the course of time, especially through the efforts of Greville (1830), Harvey (1841, 1849) and J. Agardh (1842, 1851, 1852a, 1852b, 1852e, 1863, 1872, 1876, 1879), increasing emphasis was placed on the finer details of the structure of the thallus and the reproduc- tive organs. With the appearance of J. Agardh's work of 1842, the manner of division of the tetrasporangia, whether tetrahedral, cruciate, or zonate, was also introduced into the classification of these algae. These are characters that in general are still considered important in the delimitation of taxa. As regards the characters offered by the structure of the cystocarp, J. Agardh's system, which was the standard one for some fifty years, took into account only the mature cystocarp. The multitude of significant characters presented by the ontogeny of this organ thus remained concealed, with the result that the system of J. Agardli, like those of his predecessors, contained a great deal that was artificial. The present period in the classification of these algae, which Sjostedt (1926, p. 85) has termed the embryological period, was ushered in by Schmitz's epoch- making paper of 1883. Although Nageli (1861), Bornet and Thuret (1866a, 1866b, 1867, 1876, 1878, 1880), Solms-Laubach (1867), Janczewski (1876), and Schmitz (1879b) had worked out in some detail the development of the cysto- carp, the significance of the differences in the development of this structure in different forms did not become apparent until 1883. On the basis of the funda- mental differences in the ontogeny of this organ, especially as regards the place of formation and the function of the auxiliary cell, Schmitz later (1889, 1892, and in Schmitz and Hauptfleisch, 1896-1897) proposed a regrouping of these algae along lines that portrayed a much more natural arrangement than had yet been possible. Schmitz (1892) divided the Florideophycidae into the four orders Nemalionales, Gigartinales, Rhodymeniales, and Cryptonemiales. Since comparatively few forms had been thoroughly investigated when Schmitz proposed his system, it was to be expected that further knowledge Avould necessitate revision of this system. Although Schmitz's four orders are still ac- cepted, additional developmental studies have shown that they should be re- constituted and it has been necessary to create two additional orders. The first major emendation of Schmitz's system was by Oltmanns (1904), who, among other changes, erected the order Ceramiales for those Rhod\m"ieniales of the sys- tem of Schmitz in which the auxiliary cell is formed after fertilization of the carpogonium, namely, the families Ceramiaceae, Delesseriaceae, and Rhodomela- ceae (including the Dasyaceae as currently recognized). In 1923 Kylin estab- lished the order Gelidiales for the family Gelidiaceae, which Schmitz, and fol- lowing him Oltmanns, had placed in the Nemalionales. Still later Kylin (1925) founded the order Nemastomales for the families Nemastomaceae and Rhodo- phyllidaceac (previously placed in the Cryptonemiales and Gigartinales, respec- tively) and Sjostedt (1926) erected the order Sphaerococcales for the family Sphaerococcaceae (previously placed in the Gigartinales). but these two orders were subsequently reduced by Kylin (1928, p. 113; 1932, pp. 71, 72, 76-79) under the Gigartinales. Recently Feldmann (1952) established an order Bonne- maisoniales. Although the genera comprising this order do not appear to be closely related to the other members of the Nemalionales (in which order the Bonnemaisoniaceae have been placed in recent times), the points of departure 186 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES do not seem to be of sufficient magnitude to justify recognition of a separate order. Oltmanns (1904) brought into focus an anatomical character that has proved of great importance in the classification of the Florideophycidae. He empha- sized that in some genera the thallus has a uniaxial construction whereas in others it is multiaxial. Kylin (1928, 1930a, and especially 1932) has made very effective use of this character in the separation of families. We are especially indebted to Kylin for the refinement of Schmitz's embryo- logical system of classification of the Florideophycidae. In a long series of papers, especially the monographic studies of 1923, 1928, 1930a, and 1932, he has immensely advanced our knowledge of the comparative morphology of these algae and has thereby contributed more than any other one person to a better understanding of the interrelationships of this large and diversified phy- lum. Despite certain shortcomings (see Papenfuss, 1951b) his system of 1932 allows of a much more natural arrangement than had previously been possible. It is the standard one today. In Kylin's system the orders are separated on whether or not "typical" auxiliary cells (generative auxiliary cells of Papenfuss, 1951b) are absent or present, their time of formation — before or after fertilization — and their man- ner and place of formation. Within the orders, the families are separated on whether the thallus is uniaxial or multiaxial, whether the cystocarp is imbedded in the thallus or not, whether the tetrasporangia are tetrahedrally, cruciately, or zonately divided, and various other characters of seemingly comparable importance. In regard to the long-standing disagreement between Svedelius and Kylin as to whether the Nemalionales do or do not possess a "typical" auxiliary cell reference should be made to the papers by Martin (1939), Svedelius (1942), and Kylin (1944b). In the opinion of Kylin, the cell or cells in the nemaliona- lean carpogonial branch that receive a diploid nucleus from the fertilized carpo- gonium and from which the gonimoblast develops do not constitute a "typical" auxiliary cell; yet he has no hesitation in considering the supporting cell in the Kallymeniaceae (Cryptonemiales) and in Sphaerococcus (Gigartinales) as a "typical" auxiliary cell, even though it is a cell in the carpogonial Ijranch system. Brief mention should be made of two groups of red algae which at first were not associated with this phylum. The first of these, the Corallinaceae, were for a long time, along with other calcified algae, regarded as corals. S. F. Gray (1821) appears to have been the first botanist to have considered them algae, without qualification, but they did not receive general acceptance as red algae until Decaisne (1842b) showed that they possess the typical features of this group. Despite their purple color the members of the other group, the subclass Bangiophycidae, were for many decades after they had become known classified with the filamentous or membranous green algae which they resemble in habit. As recently as 1922 Oltmanns (1922b, p. 230) stated that he was not fully con- vinced that these forms really belong with the Rhodophyceae. Although Thuret {in Le Jolis, 1863) and Rabenhorst (1868) had associated these forms with the Rhodophyceae, their place among the latter remained uncertain until the ap- pearance of Berthold's (1881a, 1882) critical investigations of various members of the group. PAPENFUSS: CLASSIFICATION OF THE ALGAE 187 Phylogeneticall}^ the red algae appear to be distantly related to the blue- green algae by way of the Bangiophycidae. Cohn (1867) was the first to arrive at this conclusion on the basis of his investigation of the pigments of both groups. Such a relationship appeared likely to Berthold (1882) also and is ac- cepted by Ishikawa (1921, 1924), Kylin (1930b, 1937a, 1943b), Tilden (1933, 1935) and Skuja (1938). In the synoptic outline that follows, Skuja's (1939a) classification of the Bangiophycidae (which has been accepted by Kylin, 1944a, and Tanaka, 1952) has been followed. The classification of the Florideophycidae is essentially that of Kylin (1932). Phylum RHODOPHYCOPHYTA Papenfuss (1946, p. 218) Syn.: Rhodophyta Wettstein (1901, p. 46) Class RiionopiiYCEAE Ruprecht (1851, p. 205) Syn.: Rhodospei-meae Harvey (1836, p. 160) ; Heterocarpeae Kiitzing (1843, p. 369) ; Pliycoerytlii'inophycees Marchand (1895, p. 17) Subclass BANGIOPHYCIDAE De Toni orth. mut. L. M. Newton (1953, p. 406) Syn.: Bangioideae De Toni (1897, p. 4) ; Protoflorideae Rosenvinge (1909, p. 55) Order PORPHYRIDIALES Kylin (1937b, p. 4; see also Kylin, 1937a, pp. 39- 51) Family Porphyridiaceae Kylin (1937b, p. 4) Order GONIOTRICHALES Skuja (1939a, p. 31) Family Goniotrichaceae (Rosenvinge) Skuja (1939a, p. 31) Family Phragmonemataceae Skuja (1939a, p. 32) Order BANGIALES Engler (1892, p. 15) Family Erythropeltidaceae Skuja (1939a, p. 33) Family Bangiaceae (S. F. Gray) Nageli (1847, p. 136) Syn.: Porphyraceae Kiitzing orth. mut. Rabenhorst (1868, p. 397); Erythrotrichiaceae Marchand orth. mut. G. M. Smith (1944, p. 162) Order RHODOCHAETALES Skuja (1939a, p. 34) Family Rhodocliaetaceae Schmitz, in Schmitz and Hauptfleisch (1896, p. 317) Order COMPSOPOGONALES Skuja (1939a, p. 34) Family Compsopogonaceae Schmitz, in Schmitz and Hauptfleisch (1896, p. 318) Subclass FLORIDEOPHYCIDAE (Lamouroux) Engler orth. mut. L. M. Newton (1953, p. 407) Syn.: Floridees Lamouroux (1813, p. 115); Euflorideae Johnson (1894, p. 639) Order NEMALIONALES Schmitz, in Engler (1892, p. 17) Syn.: Bonnemaisoniales Feldmann (1952, p. 29) Family Acrochaetiaceae Fritsch (1944, p. 258) Syn.: Chantransiaceae Kiitzing orth. mut. Rabenhorst (1868, p. 400; not including Chantransia De Candolle, see Silva, 1952, pp. 261-262); Rhodochortonaceae Nasr (1947. p. 92) Family Batrachospermaceae (C. Agardh) Dumortier orth. mut. Raben- horst (1868, p. 404) Family Lemaneaceae (S. F. Gray) Harvey orth. mut. Rabenhorst (1863, p. 275) Family Helminthocladiaceae (J. Agardh) Harvey orth. mut. Hauck (1883, p. 14) Syn.: Nemalionaceae Cohn orth. mut. G. Murray (1895, p. 207) Family Chaetangiaceae Kiitzing orth. mut. Hauck (1883, p. 14) Family Thoreaceae Reichenbach ex Hassall orth. mut. Schmitz, in Schmitz and Hauptfleisch (1896, p. 321) Family Naccariaceae Kylin (1928, p. 11) Family Bonnemaisoniaceae Schmitz, in Engler (1892, p. 20) 188 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Order GELIDIALES Kylin (1923, p. 132) Family Gelidiaceae Kiitzing orth. mut. Harvey (1853, p. 7) Order CRYPTONEMIALES Schmitz, in Engler (1892, p. 21) Family Dumontiaceae Bory orth. mut. Schmitz (1889, p. 453) Family Rhizophyllidaceae Montague orth. mut. Schmitz (1889, p. 454) Family Polyideaceae Kylin (1944a, p. 34) Syn.: Spongiocarpeae Greville (1830, p. 68) Family Squamariaceae (J. Agardh) Hauck (1883, p. 13) Family Solenoporaceae Pia (1927, p. 97) Family Hildenbrandiaceae (Trevisan) Rabenhorst (1868, p. 408) Family Corallinaceae (Lamouroux) Harvey (1849, p. 74) Family Gloiosiphoniaceae Schmitz, in Engler (1892, p. 21) Family Dermocorynidaceae Hollenberg (1940, p. 871) Family Endocladiaceae (J. Agardh) Kylin (1928, p. 41) Family Tichocarpaceae (Schmitz) Kylin (1932, p. 69) Family Cryptonemiaceae (J. Agardh) Harvey (1849, p. 75; see also Hauck, 1883, p. 16) Syn.: Grateloupiaceae Schmitz, in Engler (1892, p. 21) Family Kallymeniaceae (J. Agardh) Funk (1927, p. 389) Family Choreocolacaceae Sturch (1926, p. 602) Order GIGARTINALES Schmitz, in Engler (1892, p. 18) Syn.: Nemastomales Kylin (1925, p. 39); Sphaerococcales Sjostedt (1926, p. 75) Family Cruoriaceae (J. Agardh) Kylin (1928, p. 29) Family Calosiphoniaceae Kylin (1932, p. 5) Family Nemastomaceae (J. Agardh) Schmitz (1889, p. 453) Syn.: Gymnophlaeaceae Kiitzing (1843, p. 389); Yadranellaceae Ercegovic (1949, p. 36). The latter family is placed here at the sug- gestion of Dr. Isabella Abbott, personal communication. Family Furcellariaceae Greville orth. mut. Kylin (1932, p. 11) Family Sebdeniaceae Kylin (1932, p. 12) Family Solieriaceae (Harvey) Hauck (1883, p. 17) Family Rissoellaceae (J. Agardh) Kylin (1932, p. 31) Family Rhabdoniaceae Kylin (1925, p. 38) Family Rhodophyllidaceae Schmitz, in Engler (1892, p. 19) Family Hypneaceae J. Agardh (1852, p. 430) Family Plocamiaceae Kiitzing orth. mut. Kylin (1930, p. 45) Family Sphaerococcaceae Dumortier orth. mut. Cohn (1872a, p. 17) Family Stictosporaceae Kylin (1932, p. 53) Family Sarcodiaceae Kylin (1932, p. 54) Family Gracilariaceae (Nageli) Kylin (1930a, p. 54) Family Mychodeaceae (Schmitz et Hauptfleisch) Kylin (1932, p. 62) Family Dicranemaceae (Schmitz et Hauptfleisch) Kylin (1932, p. 65) Family Acrotylaceae Schmitz, in Engler (1892, p. 18) Family Phyllophoraceae Nageli (1847, p. 248) Family Gigartinaceae Bory orth. mut. Cohn (1880, p. 286) Family Chondriellaceae Levring (1941, p. 640) Order RHODYMENIALES Schmitz, in Engler (1892, p. 19) Family Rhodymeniaceae Harvey (1849, p. 75) Family Champiaceae Kiitzing orth. mut. Bliding (1928, p. 64) Syn.: Lomentariaceae Nageli (1847, p. 244) Order CERAMIALES Oltmanns (1904, p. 683) Family Ceramiaceae (S. F. Gray) Harvey orth. mut. Rabenhorst (1847, p. xiii) Syn.: Wrangeliaceae J. Agardh orth. mut. Harvey (1853, p. 8); Spyri- diaceae J. Agardh orth. mut. Harvey (1853, p. 8) Family Dasyaceae Kiitzing orth. mut. Rosenberg (1933, p. 83) Family Delesseriaceae Bory orth. mut. Nageli (1847, p. 208) PAPENFUSS: CLASSIFICATION OF THE ALGAE 189 Family Rhodomelaceae (J. Agardh) Harvey (1849, p. 74) Syn.: Laurenciaceae Harvey (1849, p. 74); Rytiphlaeaceae Kutzing (1843, p. 442) Prospect From the preceding review it will be evident that knowledge of the structure and reproduction of the algae, and hence of their classification, has advanced tremendously during the past hundred years. It is now well established that the assemblage of plants referred to as algae is comprised of a number of only dis- tantly related groups of organisms that share few characters except the ability of most of the forms to synthesize organic compounds by the process of photo- synthesis and the absence of a primarily produced jacket of sterile cells about the reproductive organs. If the bases of the present systems of classification of the members of the major groups are examined, however, it is found that not infrequently families and even orders have been established on the strength of knowledge obtained from a study of only a few species or in certain instances only one species. Ob- viously there exists a great need for detailed information on a large number of genera and species before it will be possible to erect schemes of classification that will portray in a reasonably accurate way the phylogenetic affinities of the organisms that constitute these groups. It is no exaggeration to say that only a good beginning has been made in the sorting out of the natural subdivisions of the major taxa. Biochemical information has contributed much to a better understanding of the interrelationships of various groups of algae. It is to be expected that biochemical investigation of the many forms that have not yet received atten- tion will yield knowledge that will be as significant as that obtained in the past. Although the chromosomes of algae are generally small and hence do not lend themselves well to karyological study, cytotaxonomic investigations like those of Cave and Pocock (1951) encouragingly point to the rich rew\ards that may be expected from the pursuit of problems in this largely unexplored area. In recent years electron microscope studies have yielded valuable informa- tion on the structure of the flagella and the cell wall of diverse algae. The inter- esting new facts brought to light augur well for an expanding use of this tool in algal research. In the past, knowledge of the developmental morphology and the life his- tories of algae has contributed greatly toward the elucidation of phylogenetic affinities among these plants. Pressing needs exist for information of this kind on many more species. In numerous instances progress in life-history studies has been greatly retarded and not infrequently the results have been woefully fragmentary owing to the difficulty of obtaining germination stages of the zygo- spores or other resting cells. It may be anticipated that in the future the physi- ology of resting cell maturation and germination will receive the attention that it merits and that the knowledge gained will make it possible to induce these cells to mature and germinate at will. An understanding of tlie physiology of resting cell germination will not only aid in life-history studies but will be a tremendous impetus to the full utilization of these simple autotrophic plants as material in experimental studies, 190 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES With the expanding use of algae as experimental material we may look for- ward to an increasing awareness among biologists in general of the need for precise identity of the forms under investigation. It may be anticipated there- fore that the preservation of voucher collections of published material will be- come standard practice and that the maintenance and welfare of such reposi- tories as herbaria, museums, and living culture collections will be the concern and the accepted responsibility, not only of the taxonomist and morphologist, but of all biologists. LITERATURE CITED Adanson, M. 1763. Families des plantes. Pt. 2. (1)-(18) + 640 + (19)-(24) pp. Paris: Vincent. Agaedh, C. a. 1817. Synopsis algarum scandinaviae . . . xl + 135 pp. Lund: Officina Berlingiana. 1824. Systema algarum. xxxviii + 312 pp. Lund: Literis Berlingianis. 1828. Species algarum . . . Vol. 2(1). Ixxvi + 189 pp. Greifswald: Moritz. 1830-1832. Conspectus criticus diatomacearum ... 66 pp. Lund: Literis Berlingianis. Agardh, J. G. 1842. Algae maris mediterranei et adriatici . . . x + 164 pp. Paris: Masson. 1848. Species genera et ordines algarum . . . Vol. 1. viii + 363 pp. Lund: Gleerup. 1851. Species genera et ordines algarum . . . Vol. 2(1). xii + 336 + [addenda and index] 337-351 pp. Lund: Gleerup. 1852a. Species genera et ordines algarum . . . Vol. 2(2:1). Pp. 337-504. Lund: Gleerup. 1852b. Species genera et ordines algarum . . . Vol. 2(2:2). Pp. 505-700 + [addenda and index] 701-720. Lund: Gleerup. 1852c. Species genera et ordines algarum . . . Vol. 2 (3:1). Pp. 701-786. Lund: Gleerup. 1863. Species genera et ordines algarum . . . Vol. 2(3:2). Pp. 787-1291. Lund: Gleerup. 1872. Bidrag till florideernes systematik. Lunds Univ. Arsskr. Afd. 2, 8(6). 60 pp. 1876. Species genera et ordines algarum . . . Vol. 3(1). Epicrisis systematis flor- idearum. vii + 724 pp. Leipzig: Weigel. 1879. Florideernes morphologi. K. Svensk. Vet.-Akad. Handl., 15(6). 199 pp., 33 pis. 1887-1888. Till algernes systematik . . . (Afd. 5). Lunds Univ. Arsskr. Afd. 3, 23(2). 174 pp., pis. 1-5. Allen, C. E. 1905. Die Keimung der Zygote bei Coleochaete. Ber. Deutsch. Bot. Ges., 23:285-292, pi. 13. Allman, G. J. 1855. Observations on ApJianizomenon Flos-a-guae, and a species of Peridinea. Quart. Journ. Micr. Sci., 3:21-25, pi. 3. 1872. Notes on Noctiluca. Quart. Journ. Micr. Sci., n. s., 12:326-332, pi. 18. Allorge, p. 1930. H^t^rocontes ou Xanthophycees? Rev. Alg., 5(2):230. Arasaki, S. 1949. On the life-history and the systematical position of Dictyosiphon foeniculaceus (Huds.) Grev. Bot. Mag. Tokyo, 62:87-90, 3 figs. PAPENFUSS: CLASSIFICATION OF THE ALGAE 191 AUESCIIOl'G, J. E. 1847. Phycearum, quae in inaribus scaudinaviae crescuut, enumeratio. Sectio prior, Fucaceas continens. Nova Acta Reg. Soc. Sci. Upsal., ser. 2, 13:223-382 + [1], pis. 1-9. Atkinson, G. P. 1905. A College Text-Book of Botany . . . , 2nd ed. xvi + 737 pp., 592 figs., frontis. New York: Holt and Co. Bary, a. de 1854. Ueber die Algengattungen Oedogonium und Bolbochaete. Ablil. Senckenberg. Naturforsch. Ges., 1:29-105, pis. 2-4. 1858a. tJber den gesclileclitlichen Zeugungsprozess bei den Algen. Ber. Verhl. Ges. Forderung Naturwiss. Freiburg i. Br., 1856:215-230, pi. 5. (Not seen, cited from Kniep, 1928.) 1858b. Untersuchungen iiber die Familie der Conjugaten (Zyguemeen und Desmid- ieen). Ein Beitrag zur physiologischen und beschriebenden Botanik. [4] + 91 pp., 8 pis. Leipzig: Forstnersche Buchhandlung. 1872. tJber den Befruchtungsvorgang bei den Charen. Monatsber. K. Preuss. Akad. Wiss. Berlin, 1871:227-239, 1 pi. 1875. Zur Keimungsgeschichte der Charen. Bot. Zeit., 33:377-385, 393-401, 409-420, pis. 5, 6. 1881. Zur Systematik der Thallophyten. Bot. Zeit, 39:1-17, 33-36. Behlau, J. 1935. Die Spondylomoraceen-Gattung Chlamydobotrys. Beitr. Biol. Pflanz., 23:125-166, 7 figs., 4 tables, pi. 3. 1939. Der Generationswechsel zwischen Chlcnnydomonas variahUis Dangeard und Carteria ovata Jacobsen. Beitr. Biol. Pflanz., 26:221-249, 3 figs. Bennett, A. W. 1878. On the structure and affinities of Characeae. Journ. Bot. 16:202-207. 1879. A few last words on Chara. Journ. Bot. 17:67-69. Bergh, R. S. 1881. Der Organismus der Cilioflagellaten. Eine phylogenetische Studie. Morph. Jahrb., 7(2) :177-288, pis. 12-16. Bergmann, C, and R. Leuckart 1852. Anatomisch-physiologische Uebersiclit des Thierreichs. xii + 690 + [2] pp., 438 figs. Stuttgart: Miiller's Verlagshandlung. Bertiiold, G. 1881a. Zur Kenntnis der Siphoneen und Bangiaceen. Mitt. Zool. Stat. Neapel, 2:72-82. 1881b. Die geschlechtliche Fortpflanzung der eigentlichen Phaeosporeen. Mitt. Zool. Stat. Neapel, 2:401-413, pi. 17. 1822. Die Bangiaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Fauna Flora Golfes Neapel, 8. [4] + 28 pp., 1 pi. BiSCHOFF, G. W. 1828. Die kryptogamischen Gewachse . . . x + 131 pp., 13 pis. Niirnberg: Schrag. Blackburn, Kathleen B. 1936. A reinvestigation of the alga Botryococcus Braunii Kiitzing. Trans. Roy. Soc. Edinburgh, 58(3) : 841-854, 4 figs., pi. 1. Blackman, F. F. 1900. The primitive algae and the Flagellata. An account of modern work bearing on the evolution of the algae. Ann. Bot., 14:647-688, figs. 13, 14. 192 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Blackman, F. F., and A. G. Tansley 1902. A revision of tlie classification of the green algae. New Phytol., 1:17-24, 47-48, 67-72, 89-96, 114-120, 133-144, 163-168, 189-192, 213-220, 238-244. Eliding, C. 1928. Studien iiber die Florideenordnung Rhodymeniales. Lunds Univ. Arsskr., N. F., Avd. 2, 24(3). 74 pp., 52 figs. BOHLIN, K. 1897a. Studier ofver nagra slagten af alggruppen Confervales Borzi. Bihang K. Svensk. Vet.-Akad. Handl. 23, Afd. 3, No. 3. 56 pp., 2 pis. 1897b. Zur Morphologie und Biologie einzelliger Algen. (Vorlaufige Mitteilung.) ofvers. K. [Svensk.] Vet.-Akad. Forhandl. 1897:507-529, 10 figs. 1901. Utkast till de grona algernas och arkegoniaternas fylogeni. 43 + iv pp., 1 fold- ing chart. Thesis. Lund: Publ. by author. BONMEMAISON, T. 1822. Essai d'une classification des hydrophites loculees, ou plantes marines articu- lees qui croissant en France. Journ. Phys., Chimie, Hist. Nat. et Arts, 94:138-148 (Mars), 174-203 (Avril). BoNPLAND, A., and F. H. A. von Humboldt 1815. Nova genera et species plantarum . . . Tome 1. Iviii + 377 pp., 96 pis. Paris: Publ. by authors? BORGE, 0., and A. Pascher 1913. Zygnemales. In Pascher, Die Susswasser-Flora Deutschlands, osterreichs und der Schweiz, Heft 9. iv + 51 pp., 79 figs. Jena: G. Fischer. BORGERT, A. 1891. tJber die Dictyochiden, insbesondere iiber Distephanus speculum; sowie Studien an Phaeodarien. Zeitschr. Wiss. Zool., 51:629-676, 2 figs., pi. 33. B0RGESEN, F. 1905. Contributions a la connaissance du genre Siphonocladus Schmitz. Overs. K. Dansk. Vidensk. Selsk. Forhandl., 1905:259-291, 13 figs. 1913. The marine algae of the Danish West Indies. Part 1. Chlorophyceae. Dansk Bot. Arkiv 1(4). 158 + [2] pp., 126 figs., 2 maps. 1925. Marine algae from the Canary Islands ... I. Chlorophyceae. K. Dansk. Vidensk. Selsk. Biol. Meddel., 5(3). 123 pp., 49 figs. BORNET, £}. 1891. Note sur quelques Ectocarpus. Bull. Soc. Bot. France, 38:353-372, pis. 6-8. BoBNET, £;., and C. Flahault 1886-1888. Revision des Nostocac^es heterocystees contenues dans les principaux herbiers de France. Ann. Sci. Nat. Bot., ser 7, 3:323-381 (1886); 4:343-373 (1886); 5:51-129 (1887); 7:177-262 (1888). BoRNET, £;., and G. Thuret 1866a. Sur la fecondation des Floridees. Compt. Rend. Acad. Sci. [Paris], 63:444-447. 1866b. Note sur la fecondation des Floridees. Mem. Soc. Imp. Sci. Nat. Cherbourg, 12:257-262. 1867. Recherches sur la fecondation des Floridees. Ann. Sci. Nat. Bot., ser. 5, 7:137- 166, pis. 11-13. 1876. Notes algologiques . . . , fasc. 1. xx + 1-70 + [2] pp., pis. 1-25. Paris: Masson. 1880. Notes algologiques . . . , fasc. 2. Pp. 71-196, pis. 26-50. Paris: Masson. PAPENFUSS: CLASSIFICATION OF THE ALGAE 193 BORZI, A, 1878. Note alia inorfologia e biologia delle alghe ficocromacee. Nuova Giorn. Bot. Ital., 10:236-288, pis. 7-10. 1879. Note alia morfologia e biologia delle alghe ficocromacee. Nuovo Giorn. Bot. Ital., 11:347-388, pis. 9-12. 1882. Note alia morfologia e biologia delle alghe ficocromacee. Nuovo Giorn. Bot. Ital., 14:272-315, pis. 16, 17. 1889. Botrydiopsis nuovo genere di alghe verdi. Boll. Soc. Ital. Microscop., 1:60-70. 1895. Studi algologici . . . , fasc. 2. Pp. [i-vii], 121-378, pis. 10-31. Palermo: Reber. 1914. Studi sulle mixoficee. Nuovo Giorn. Bot. Ital., ser. 2, 21:307-360. 1916. Studi sulle mixoficee. Nuovo Giorn. Bot. Ital., ser. 2, 23:559-588, 6 figs. 1917. Studi sulle mixoficee. Nuovo Giorn. Bot. Ital., ser. 2, 24:17-30, 65-112, 198-214, pis. 6-10. Braun, a. 1849. Uebersicht der schweizerischen Characeen. Ein Beitrag zur Flora der Schweiz. Neue Denkschr. Allg. Schweiz. Ges. Gesammt. Naturwiss., 10 (Art. 4). 23 pp. 1851. Betrachtungen fiber die Erscheinung der Verjungung in der Natur, insbeson- dere in der Lebens- und Bildungs-geschichte der Pflanze. xvi + 363 pp., 3 pis. Leipzig: W. Engelmann. 1852. tJber die Richtungsverhaltnisse der Saftstrome in den Zellen der Characeen. Ber. Verb. K. Preuss. Akad. Wiss. Berlin, 1852:220-268. 1853. iJber die Richtungsverhaltnisse der Saftstrome in dev Zellen der Characeen (Fortsetzung und Schluss). Ber. Verh. K. Preuss. Akad. Wiss. Berlin, 1853: 45-76. 1855. Algarum unicellularium genei'a nova et minus cognita. [6] + 111 pp., 6 pis. Leipzig: W. Engelmann. Beaun, a., and 0. Nordstedt 1883. Fragmente einer Monographie der Characeen. Abh. K. Akad. Wiss. Berlin, 1882 (Article 1). 211 pp., 7 pis. Brown, H. P. 1945. On the structure and mechanics of the protozoan fiagellum. Ohio Journ. Sci., 45:247-278, pis. 1-12. BRUN NTH ALEE, J. 1913. Die systematische Gliederung der Protococcales (Chlorophyceae). Verh. Zool.- Bot. Ges. Wien, 63:76-91. 1915. Protococcales. In A. Pascher, Die Susswasser-Flora Deutschlands, osterreichs und der Schweiz, Heft 5: Chlorophyceae II. Pp. 52-205, 330 figs. Jena: G. Fischer. BXJTSCHLI, O. 1883-1887. Protozoa. Abt. 2, Mastigophora. In H. G. Bronn's Klassen und Ordnungen des Thier-Reichs . . . Bd. 1. Pp. [3], 617-1097, pis. 39-55. Leipzig and Heidelberg: Winter'sche Verlagshandlung. 1885. Einige Bemerkungen fiber gewisse Organisations-verhaltnisse der sog. Cilio- flagellaten und der Noctiluca. Morph. Jahrb. 10:529-577, 4 figs., pis. 26-28. Calvin, M., and V[ictoria] Lynch 1952. Grana-like structures of Synechococcus cedorum. Nature, 169:455-456, 2 figs. Carter, H. J. 1856. Notes on the freshwater Infusoria of the Island of Bombay. No. 1. Organiza- tion. Ann. Mag. Nat. Hist., ser. 2, 18:115-132, 221-249, pis. 5-7. 1858. Note on the red colouring matter of the sea 'round the shores of the Island of Bombay. Ann. Mag. Nat. Hist., ser. 3, 1:258-262. 194 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Carteb, Nellie 1937. New or interesting algae from brackish water. Archiv Protistenk. 90(1): 1-68, 3 figs., pis. 1-8. Cave, Marion S., and Mary A. Pocock 1951. Karylogical studies in the Volvocaceae. Amer. Journ. Bot, 38:800-811, 47 figs., 2 tables. Chadefaud, M. 1945. Sur la position systematique des Vaucheries. Bull. Soc. Bot. France, 92:47-50. 1950a. Les cellules nageuses des algues dans rembranchement des Chromophycees. Compt. Rend. Acad. Sci. [Paris], 231:788-790, 7 figs. 1950b. Les cellules nageuses des algues dans rembranchement des Chlorophyc^es. Compt. Rend. Acad. Sci. [Paris], 231:988-990, 7 figs. Chapman, V. J. 1952. New entities in the Chlorophyceae of New Zealand. Trans. Roy. Soc. New Zealand, 80:47-58, 12 figs., pis. 21, 22. Chatton, IE 1920. Les p^ridiniens parasites morphologie, reproduction, ethologie. Arch. Zool. Experiment. Gen., 59:1-475, 161 figs., 18 pis. C HOD AT, R. 1902. Algues vertes de la Suisse. Pleurococcoides-Chrool^poides. Beitr. Krypt.-flora Schweiz, 1(3). xiii + 373 pp., 264 figs. 1909. Etude critique et experimentale sur le polymorphisme des algues. 165 + [1] pp., pis. 1-21, A-B. Geneva: Georg and Co. 1921. Materiaux pour I'histoire des algues de la Suisse. IV. Sur le groupe provisoire des Chrysostomatacees. Bull. Soc. Bot. Geneve, ser. 2, 13:81-87, figs. 8-10. Cholnoky, B. von 1928. tJber die Auxosporenbildung der Anomoeoneis sculpta E. CI. Archiv Protistenk., 63:23-57, pis. 3-6. 1933a. Beitrage zur Kenntnis der Karyologie der Diatomeen. Archiv Protistenk., 80:321-348, 41 figs. 1933b. Die Kernteilung von Melosira arenaria nebst einigen Bemerkungen iiber ihre Auxosporenbildung. Zeitschr. Zellforsch. Mikr. Anat., 19:698-719, 24 figs. Church, A. H. 1919a. Historical review of the Phaeophyceae. Journ. Bot., 57:265-273. 1919b. Historical review of the Florideae.— I. Journ. Bot., 57:297-304. [The same]— II. lUd.: 329-334. ClENKOWSKI, L. 1865a. Ueber einige chlorophyllhaltige Gloeocapsen. Bot. Zeit., 23:21-27, pi. 1. 1865b. Beitrage zur Kenntniss der Monaden. Archiv Mikr. Anat., 1:203-232, pis. 12-14. 1870. Ueber Palmellaceen und einige Flagellaten. Archiv Miki\ Anat., 6:421-438, pis. 23, 24. Claparede, E., and J. Lachmann 1858-1859. :fitudes sur les infusoires et les Rhizopodes. "Vol. 1. Mem. Inst. Genevois, Vols. 5 and 6. 482 pp., 24 pis. Cleland, R. E. 1919. The cytology and life history of Nemalion multifidum, Ag. Ann. Bot., 33:323- 351, 3 figs., pis. 22-24. PAPENFUSS: CLASSIFICATION OF THE ALGAE 195 COHN, P. 1852. Ueber eine neue Gattung aus der Familie der Volvocinen. Zeitschr. Wiss. Zool., 4(1): 77-116, pi. 6. 1853. Beitrage zur Entwickelungsgeschichte der lufusorien. II. Ueber den Encysti- rungsprocess der lufusorien. Zeitschr. Wiss. Zool., 4(3-4) :253-281, pi. 13. 1854. Untersuchungen iiber die Entwickelungsgeschichte der mikroskopischen Algen und Pilze. Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Curios., 24:103-256, pis. 15-20. 1855. tJber die Fortpflanzung von Sphaeroplea annuUna. Ber. Verb. K. Preuss. Akad. Wiss. Berlin, 1855:335-351. 1856. Observations sur les Volvocinees, et specialement sur I'organisation et la propagation du Volvox glohator. Ann. Sci. Nat. Bot., ser. 4, 5:323-332. 1867. Beitrage zur Physiologic der Phycochromaceen und Florideen. Archiv Mikr. Anat, 3:1-60, pis. 1, 2. 1872a. Conspectus familiarum cryptogamarum secundum metbodum naturalem dis- positarum. Hedwigia, 11:17-20. 1872b. Grundziige einer neuen natiirlichen Anordnung der kryptogamischen Pflanzen. Jahres-Ber. Schles. Ges. Vaterl. Cultur, 49:83-89. 1880. tJber sein Thallophytensystem. Jahres-Ber. Schles. Ges. Vaterl. Cultur, 57: 279-289. CONEAD, W. 1914. Contributions a I'etude des flagellates: III. La niorphologie et la nature des enveloppes chez Hymenomonas roseola Stein et H. coccolithophora Massart et Conrad, nov. spec, et les Coccolithophoridae. Ann. Biol. Lacustre, 7:155- 164, 6 figs. 1926. Recherches sur les Flagellates de nos eaux saumatres. 29 Partie: Chrysomona- dines. Archiv Protistenk., 56:167-231, 28 figs., pis. 7-9. 1927. Essai d'une monographie des genres Mallomonas Perty (1852) et Pseudo- mallomonas Chodat (1920). Archiv Protistenk., 59:423-505, 42 figs., pis. 8-11. 1928. Le genre Microglena C. G. Ehrenberg (1838). Archiv Protistenk., 60:415-439, 13 figs. 1933. Revision du genre Mallomonas Perty (1851) incl. Pseudomallomonas Chodat (1920). Mem. Mus. Roy. Hist. Nat. Belgique, No. 56. 82 pp., 70 figs. COPELAND, J. J. 1936. Yellowstone thermal Myxophyceae. Ann. New York Acad. Sci., 36:i-vi, 1-229, 73 figs. CZURDA, V. 1932. Zygnemales [second treatment]. In A. Pascher, Die Siisswasser-Plora Mittel- europas. Heft 9. iv + [1] +232 pp., 226 figs. Jena: G. Fischer. 1937. Conjugatae. In K. Linsbauer, Handbuch der Pflanzenanatomie. Abt. II, Bd. 6(2) : Algen B:b. [2] + 176 pp., 124 figs. Berlin: Gebr. Borntraeger. Dalechamps, J. 1587. Historia generalis plantarum, . . . Vol. 1. [10] + 1095 pp., illus. with unnum- bered text figs. Lyons: Rovillius. Dangeard, p. 1937. Notice sur la vie et les travaux de Camille Sauvageau (1861-1936). Bull. Stat Biol. Arcachon, 34:5-57, portr. Dangeard, P. A. 1889. Recherches sur les Cryptomonadinae et les Euglenae. Botaniste, 1:1-38, pi. 1. 196 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Decaisne, J. 1842a. Essai sur une classification des algues et des polypiers calciferes de Lamouroux. Ann. Sci. Nat. Bot, s^r. 2, 17:297-380, pis. 14-17. 1842b. Memoire sur les corallines ou polypiers calciferes. Ann. Sci. Nat. Bot., ser. 2, 18:96-128. Deflandre, G. 1934. Sur la structure des flagelles. Ann. Protistol., 4:31-54, pis. 1-5. Derbes, a., and A. J. J. Solier 1850. Sur les organes reproducteurs des algues. Ann. Sci. Nat. Bot., ser. 3, 14:261- 282, pis. 32-37. Desikachary, T. V. 1952. Electron microscope study of the diatom-wall structure. Journ. Sci. Indus. Res. llB(ll): 491-500, 28 figs. DiESING, K. M. 1866. Revision der Prothelminthen. Abtheilung: Mastigophoren. Sitzungsber. Math.- Naturwiss. CI. K. Akad. Wiss. Wien, 52 Abt. 1:287-401. DiLLWYN, L. W. 1802-1809. British Confervae; or Colored Figures and Descriptions of the British Plants Referred by Botanists to the Genus Conferva. [2] + 87 + [231] pp., pis. 1-109, A-G. London: Phillips. DlWALD, K. 1938. Die ungeschlechtliche und geschlechtliche Fortpflanzung von Glenodinium luMniensiforme spec. nov. Flora, 132:174-192, 4 figs., 5 tables. DOFLEIN, F. 1922. Untersuchungen iiber Chrysomonadinen. Archiv Protistenk., 44(2) : 149-213, 4 figs., pis. 6-10. 1923. Untersuchungen fiber Chrysomonaden. Ill and IV. Archiv Protistenk., 46: 267-344, 7 figs., pis. 15-22. 1928. Lehrbuch der Protozoenkunde . . . 5th ed. (E. Reichenow, ed.), Teil 2, Spezielle Naturgeschichte der Protozoen, Erste Halfte: Mastigophoren und Rhizopoden. Pp. 439-862, figs. 389-823. Jena: G. Fischer. Drew, Kathleen M. 1944. Nuclear and somatic phases in the Florideae. Biol. Rev., 19:105-120, 6 figs. Droiet, F. 1951. Cyanophyta. In G. M. Smith, ed., Manual of Phycology ... Pp. 159-166. Waltham, Mass.: Chronica Botanica Co. Drouet, F., and A. Cohen 1935. The morphology of Gonyostomum semen from Woods Hole, Massachusetts. Biol. Bull., 68:422-439, 2 pis. 1937. Further observations on Gonyostomum semen. Bot. Gaz., 98:617-618, 1 fig. Drouet, F., and W. A. Daily 1952. A synopsis of the coccoid Myxophyceae. Butler Univ. Bot. Studies, 10:220-223. Dujardin, F. 1841. Histoire naturelle des zoophytes. Infusoires, . . . xii + 684 pp. + atlas of 14 + [2] pp. and 22 pis. Paris: De Boret. DUMORTIER, B. C. 1822. Commentationes botanicae. 116 + [1] pp., 1 folding leaf. Tournay: Ch. Caster- man-Dieu. PAPENFUSS: CLASSIFICATION OF THE ALGAE 197 DUMORTIEK, B. C. (CoUt.) 1829. Analyse des families des plantes, avec I'lndication des principaux genres qui s'y rattachent. 104 pp. Tournay: J. Casterman. Egerod, Lois E. 1952. An analysis of the siphonous Chlorophycophyta with special reference to the Siphonocladales, Siphonales and Dasycladales of Hawaii. Univ. Calif. Publ. Bot., 25:325-454, 23 figs., pis. 29-42. Ehrenberg, D. C. G. 1832. Tiber die Entwickelung und Lebensdauer der Infusionsthiere; nebst ferneren Beitragen zu einer Vergleichung ihrer organischen Systeme. Abh. K. Akad. Wiss. Berlin, 1831:1-154, 4 pis. 1836. Bemerkungen iiber feste mikroskopische, anorganische Formen in den erdigen und derben Mineralien. Ber. Verb. K. Preuss. Akad. Wiss. Berlin, 1836:84-85. 1838. Die Infusionsthierchen als vollkommene Organismen. Ein Blick in das tiefere organische Leben der Natur. xviii + [4] + 547 + [1] pp. and atlas of 64 pis. Leipzig: L. Voss. 1839. tJber die Bildung der Kreidefelsen und des Kreidemergels durch unsichtbare Organismen. Abh. K. Akad. Wiss. Berlin, 1838:59-147, 3 charts, 4 pis. 1841. tJber noch jetzt zahlreich lebende Thierarten der Kreidebildung und den Or- ganismus der Polythalamien. Abh. K. Akad. Wiss. Berlin, 1939:81-174, 4 pis. 1853. Neuerlich bei Berlin vorgekommenen neuen Formen des mikroskopischen Lebens. Monatsber. Berlin Akad. Wiss., 1853:183-194. ElCHLER, A. W. 1880. Syllabus der Vorlesungen iiber specielle und medicinisch-pharmaceutische Botanik. iv + 47 pp. Berlin: Gebr. Borntraeger. Elenkin, a. a. 1916. Note sur le genre Nodularia Mert., suivie de la recherche critique sur la famille Nodulariaceae mihi. Bull. Jardin Imp. Bot. Pierre le Grand, 16(2) : 333-344. (Not seen, cited from Elenkin, 1917, p. 163.) 1917. M^moire sur la modification des principes de classification des Hormogoneae (Thur.) Kirchn. (tribu des Cyanophyc^es). Journ. Soc. Bot. Russie, 1(3-4): 147-165, 5 figs. 1933. tJber die systematische Einteilung der Ordnung Chroococcales Geitler (1925). Akad. Nauk Bot. Inst. Trudy (Acta Inst. Bot. Acad. Sci. U.S.S.R.), ser. 2, PI. Crypt., Fasc. 1:17-21. (In Russian with a German summary.) 1934. Sur les principes fondamentaux du systeme des Cyanophyceae. Soviet Bot., 1934(5) :51-83. (In Russian.) 1936. Monographia algarum cyanophycearum aquidulcium et terrestrium in finibus URSS inventarum. Vol. 1. 675 pp. Moscow, Leningrad. (Not seen, cited from Geitler, 1942.) 1938. Monographia algarum cyanophycearum aquidulcium et terrestrium in finibus URSS inventarum. Pars specialis (systematica) . . . Fasc. I . . . 984 pp., 290 figs. Moscow, Leningrad: Acad. Nauk. 1949. Monographia algarum cyanophycearum aquidulcium et terrestrium in finibus URSS inventarum. Pars specialis (systematica). Fasc. II . . . Pp. 985-1908, figs. 291-566, portr. Moscow, Leningrad: Acad. Nauk. Ellis, J. 1767. Sequel [to] extract of a letter from John Ellis, Esquire, F.R.S. to Dr. Linnaeus, of Upsal[a], F.R.S. on the animal nature of the genus of zoophytes, called Corallina. Philos. Trans. [Roy. Soc. London], 57:421-427, pi. 18. 198 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Endlicher, S. L. 1836-1840. Genera plantarum secundum ordines naturales disposita. Ix + 1483 pp. Vienna: Beck. 1843. Mantissa botanica altera. Sistens generum plantarum, suppl. 3. [6] + 111 pp. Vienna: Beck. Engler, a. 1892. Syllabus der Vorlesungen iiber specialle und medicinisch-pharmaceutische Botanik. Eine Uebersicht iiber das gesammte Pflanzensystem mit Beriick- sichtigung der Medicinae- und Nutzpflanzen. Grosse Ausg. xxiii + 184 pp. Berlin: Gebr. Borntraeger. 1897. Bemerkung, betreffend der in der Abteilung I. 2 noch niclit beriicksichtigten Chlorophyceae und Phaeophyceae. In A. Engler and K. Prautl, Die natiir- lichen Pflanzenfamilien . . . Teil 1, Abt. 2, p. 570. Leipzig: W. Engelmann. 1898. Syllabus der Pflanzenfamilien . . . 2nd ed. xii + 214 pp. Berlin: Gebr. Born- traeger. 1903. Syllabus der Pflanzenfamilien . . . 3rd ed. xxvii + 233 pp. Berlin: Gebr. Born- traeger. Engler, A., and E. Gilg 1924. Syllabus der Pflanzenfamilien . . . 9tli and lOth eds. xliii + 420 pp. Berlin: Gebr. Borntraeger. Ercegovic, a. 1932. Ekoloske i socioloske studije o litofitskim cijanoficejama sa jugoslavenske obale Jadrana. Rad Jugoslav. Akad. Znanosti Umjetnosti, 244:129-220, 10 figs., 11 tables, 7 pis. 1949. Sur la Yadranella, nouveau genre d'algues de I'Adriatique et sur son develop- pement. Acta Adriatica Inst. Oceanogr. Ribarstvo Split, 4(2) : 27-39, 4 figs. ESPER, E. J. C. 1797-1808. Icones fucorum ... 217 + 53 + 53bis-132 +iv + iii pp., pis. 1-29, 29A, 30-136, 136A, 137-144, 144A, 145-160, 160A, 161, 161A, 162, 162A, 163, 163A, 164-169, I-IV, I-IV. Niirnberg: Raspeschen Buchhandlung. Falkenberg, p. 1879. Die Befruchtung und der Generationswechsel von Cutleria. Mitt. Zool. Stat. Neapel, 1:420-447, pi. 13. 1882. Die Algen in weitesten Sinne. Zn A. Sclienk, Handbuch der Botanik. Pp. 159-314, 25 figs. Breslau: Eduard Trewendt. Farmer, J. B., and J. L. Wiliams 1896. On fertilization and the segmentation of the spore in Fucus. Ann. Bot., 10: 479-487. 1898. Contributions to our knowledge of the Fucaceae: their life-history and cytology. Philos. Trans. Roy. Soc. London, ser. B, 190:623-645, pis. 19-24. Feldmann, J. 1937. Les algues marines de la cote des Alberes. I-III. Cyanophycees, Chlorophycees, Pheophycees. Rev. Alg., 9(3-4): 141-335, 67 figs., 10 pis. 1938. Sur la classification de I'ordre des Siphonocladales. Rev. Gen. Bot., 50:571-597, 5 figs. 1946. Sur I'heteroplastie de certaines Siphonales et leur classification. Compt. Rend. Acad. Sci. [Paris], 222:752-753. 1949. L'ordre des Scytosiphonales. Mem. Hist. Nat. Afr. Nord, hors-ser., 2:103-115, 3 figs. PAPENFUSS: CLASSIFICATION OF THE ALGAE 199 Feldmann, J. (Cont.) 1950. Sur I'existence d'une alternance de generations entre VHalicystis parvula Schmitz et le Derhesia tenuissima (De Not.) Crn. Compt. Rend. Acad. Sci. [Paris], 230:322-323. 1952. Les cycles de reproduction des algues et leur rapports avec la phylog^nie. Rev. Cytol. Biol. Veg., 13:1-49. FORTI, A. 1907. Myxophyceae. In J. B. De Toni, Sylloge algarum . . . Vol. 5 [6] + 761 pp. Padua: Publ. by De Toni. FOSLIE, M. 1890. Contributions to knowledge of the marine algae of Norway. I. East-Finmarken. Troms0 Mus. Aarshefter, 13:1-186, pis. 1-3. F0YN, BiRGITHE R. 1934. tJber den Lebenscyklus einiger Braunalgen. Vorlaufige Mitteilung. Bergens Mus. Arbok, 1934:1-9. F0YN, Bjorn 1929. Untersuchungen iiber die Sexualitat und Entwicklung von Algen. IV. Vorlau- fige Mitteilung iiber die Sexualitat und den Generationswechsel von Clado- phora und Ulva. Ber. Deutsch. Bot. Ges., 47:495-506, 2 figs., 2 tables. 1934a. Lebenszjidus, Cytologic und Sexualitat der Chlorophycee Cladophora Suhriana Kiitzing. Archiv Protistenk., 83:1-56, 18 figs., 13 tables, pis. 1-5. 1934b. Lebenszyklus und Sexualitat der Chlorophycee Ulva lactuca L. Archiv Protis- tenk., 83:154-177, 13 figs., 8 tables. Fremy, p. 1930. Les myxophycees de I'Afrique equatoriale frangaise. Arch. Bot., 3 (Mem. 2). 508 pp., 362 figs., 1 folding map. 1933. Cyanophycees des cotes d'Europe. Mem. Soc. Natl. Sci. Nat. et Math. Cherbourg, 41:1-235, pis. 1-66. Fritsch, F. E. 1929. The genus Sphaeroplea. Ann. Bot, 43:1-26, 8 figs. 1935. The Structure and Reproduction of the Algae, Vol. 1. xvii + 791 pp., 245 figs., frontis. Cambridge Univ. Press. 1939. The heterotrichous habit. Bot. Not., 1939:125-133. 1942. The interrelations and classification of the Myxophyceae (Cyanophyceae). New Phytol., 41:134-148, 5 figs. 1943. Studies in the comparative morphology of the algae III. Evolutionary tenden- cies and affinities among Phaeophyceae. Ann. Bot., n.s., 7:63-87, 4 figs. 1944. Present-day classification of algae. Bot. Rev., 10:233-277. 1945. The Structure and Reproduction of the Algae. Vol. 2. xiv -f 939 pp., 336 figs., 2 maps, frontis. Cambridge Univ. Press. 1947. The status of the Siphonocladales. Journ. Indian Bot. Soc, M.O.P. Iyengar Commemoration Volume, pp. 29-50, 5 figs. 1951. The heterocyst: A botanical enigma. Proc. Linn. Soc. London, 162(2) :194-211, 122 figs., pi. 5. Fujiyama, T. 1949. On the asexual reproduction and life-history of Prasiola (Ag.) Menegh. in Japan. Bot. Mag. Tokyo, 62:57-61, 10 figs. Funk, G. 1927. Die Algenvegetation des Golfs von Neapel . . . Publ. Staz. Zool. Napoli, 7 (suppl.). 507 pp., 50 figs., 20 pis. 200 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Geitlee, L. 1925a. Synoptische Darstellung der Cyanophyceen in morphologischer und systemati- scher Hinsicht. Beih. Bot. Cent., 41(3) : 163-294, pis. 17-20. 1925b. Cyanophyceae. In A. Pascher, Die Siisswasser-Flora Deutschlands, ossterreichs und der Schweiz. Heft 12. viii + 450 + 464-481 pp., 560 figs. Jena: G. Fischer. 1927a. Die Reduktionsteilung und Copulation von Cymhella lanceolata. Archiv Pro- tistenk., 58:465-507, 14 figs., pis. 8, 9. 1927b. Somatische Teilung, Reduktionsteilung, Copulation und Parthenogenese bei Cocconeis placentula. Archiv Protistenk., 59:506-549, 29 figs., pis. 12-14. 1928. Copulation und Geschlechtsverteilung bei einer Nitzschia-Art. Archiv Protis- tenk., 61:419-442, 13 figs. 1930-1932. Cyanophyceae. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2nd ed. Bd. 14 (issued by R. Kolkwitz). vi + 1196 pp., 780 figs. Leipzig: Akad. Verlagsges. 1933. Diagnosen neuer Blaualgen von den Sunda-Inseln. Arch. Hydrobiol. Suppl., 12:622-634. 1935. Trypayiochloris, eine neue griine Alge in den Schalen von Landschnecken und ihre Begleitfiora. Biol. Generalis, 11(2) : 135-148, 5 figs., pis. 2, 3. 1942. Schizophyta: Klasse Schizophyceae. In A. Engler and K. Prantl, Die Natiir- lichen Pfianzenfamilien . . . 2nd ed. (cont. by H. Harms and J. Mattfeld). Bd. lb. iv + 232 pp., 156 figs. Leipzig: W. Engelmann. 1943. Eine neue atmophytische Chrysophycee, Ruttnera spectabilis, nov. gen., nova spec. Internat. Rev. Ges. Hydrobiol. Hydrograph. 43(1-3): 100-109, 2 figs. 1952. Oogamie, Mitose, Meiose und metagama Teilung bei der zentrischen Diatomee Cyclotella. osterr. Bot. Zeitschr., 99:506-520, 5 figs. Gemeinhakdt, K. 1930. Silicoflagellatae. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2nd ed. Bd. 10, Abt. 2 (issued by R. Kolkwitz). Pp. 1-85, 69 figs., pi. 1. Leipzig: Akad. Verlagsges. 1931. Die Silicoflagellaten der Deutschen Sudpolar-Expedition 1901-1903. In E. von Drygalski, Deutsche Sudpolar-Expedition 1901-1903 im Auftrage des Reichs- ministeriums des Innern, Bd. 20, pp. 217-258, 1 fig., pis. 42, 43. Berlin and Leipzig: Walter De Gruyter and Co. 1938-1940. Oedogoniales. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2nd ed. Bd. 12, Abt. 4 (issued by R. Kolkwitz). xii + 453 pp., 539 figs. Leipzig: Akad. Verlagsges. Gmelin, S. G. 1768. Historia fucorum. [10] +239 + 6 pp., pis. lA, IB, 2A, 2B, 3-33. Leningrad: Acad. Sci. 1788. See Linnaeus, C. 1788. Goebel, K. 1930. Die Deutung der Characeen-Antheridium. Ein Versuch. Flora, 124:491-498, 3 figs. GOLDFUSS, G. A. 1820. Handbuch der Zoologie. Vol. 1. xlvi + 696 pp., 2 pis. Niirnberg: Schrag. GOMONT, M. 1892. Monographia des oscillari^es (nostocac^es homocyst^es). Ann. Sci. Nat. Bot., s^r. 7, 15:263-368, pis. 6-14. 1893. Monographia des oscillariees (nostocac^es homocyst^es) deuxieme partie- lyngbyees. Ann. Sci. Nat. Bot., s^r. 7, 16:91-264, pis. 1-7. PAPENFUSS: CLASSIFICATION OF THE ALGAE 201 GOROSCHANKIN, J. 1891. Beitrage zur Kenntniss der Morphologie und Systematik der Chlamydomonaden. I. Chlamydomonas Braunii (Mihi). Bull. Soc. Imp. Nat. Moscou, n. s., 4:498- 518, pis. 14, 15. Gkaham, H. W. 1951. Pyrrophyta. In G. M. Smith, ed., Manual of Phycology ... Pp. 105-118, figs. 26-28. Waltham, Mass.: Chronica Botanica Co. Gray, S. F. 1821. A Natural Arrangement of British Plants, . . . Vol. 1. xxviii + 824 pp., 21 pis. London: Baldwin, Cradock, and Joy. Greville, R. K. 1830. Algae britannicae, . . . Ixxxviii + 218 pp., 19 pis. Edinburgh: Maclachlan and Stewart. Gkiffeth, J. W., and A. Henfrey 1856. The Micrographic Dictionary; a Guide to the Examination and Investigation of the Structure and Nature of Microscopic Objects, xl + 696 pp., 816 figs., 41 pis. London: van Voorst. Gross, F. 1934. Zur Biologie und Entwicklungsgeschichte von Noctiluca miliaris. Archiv Pro- tistenk., 83:178-196, 6 figs., pi. 6. Groves, J., and G. R. Bullock-Webster 1920. The British Charophyta. Vol. 1. xiv + 141 pp., pis. 1-20. London: Ray Society. 1924. The British Charophyta. Vol. 2. xi + 129 pp., pis. 21-45. London: Ray Society. Gbunow, A. 1860. Ueber neue Oder ungenugend gekannte Algen. Verb. Zool.-Bot. Ges. Wien, 10: 503-582, pis. 3-7. Haeckel, E. 1862. Die Radiolarien. (Rhizopoda Radiaria.) Eine Monographie, Thiel 1. xiv + 572 pp. Berlin: G. Reimer. 1866. Generelle Morphologie der Organismen. Bd. 1. Allgemeine Anatomie der Organismen. xxxii + 574 pp., 2 pis. Berlin: G. Reimer. 1873. Naturliche Schopfungsgeschichte. 4th ed. xlvi + 688 pp., 16 pis. Berlin: G. Reimer. (Not seen, cited from Kofoid, in Univ. Calif. Publ. Zool., 19:329, 1920.) 1894. Systematische Phylogenie der Protisten und Pflanzen. Theil 1. xv + 400 pp. Berlin: G. Reimer. Hamel, G. 1931-1939. Pheophyc^es de France. 1-80 + i-xlvii + 81-432 pp., 63 figs., 10 pis. Paris: Publ. by author. Hammerling, J. 1931. Entwicklung und Formbildungsvermogen von Acetabularia niediterranea. Biol. Zentralbl., 51:633-647, 6 figs., 1 table. Hansgirg, a. 1886. Prodromus der Algenflora von Bohmen, Theil 1, Heft 1. Archiv Naturwiss. Landesdurchforsch. Bohmen, 5, No. 6. 96 pp., 45 figs. 1888a. Prodromus der Algenflora von Bohmen, Theil 1, Heft 2. Archiv Naturwiss. Landesdurchforsch. Bohmen, 6, No. 6. 288 + [2] pp., figs. 46-124. 202 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Hansgikg, a. (Cont.) 1888b. Synopsis generum subgenerumque myxophycearum (cyanophycearum) hucus- que cognitorum cum descriptione generis nov. "Dactylococcopsis." Notarisia, 3:548-590. 1892. Prodromus der Algenflora von Bohmen. Tlieil 2. Archiv Naturwiss. Landesdurch- forsch. Bohmen, 8, No. 4. 268 pp., 68 figs. Hartmann, M. 1929. Untersuchungen iiber die Sexualitat und Entwicklung von Algen. III. tJber die Sexualitat und den Generationswechsel von Chaetomorplia und Entero- morpha. Ber. Deutsch. Bot. Ges., 47:485-494, 1 fig., 2 tables. Harvey, W. H. 1836. Algae. In J. T. Mackay, Flora hibernica . . . , Pt. 3. Pp. 157-254. Dublin: William Curry and Co. 1841. A Manual of the British Algae . . . Ivii + 229 pp. London: van Voorst. 1847. Nereis australis . . . viii + 124 pp., 50 pis. London: Reeve Bros. 1849. A Manual of the British Marine Algae . . . 2nd ed. lii + 252 pp., 27 pis. London: van Voorst. 1852. Nereis boreali-americana: . . . Pt. 1. — Melanospermeae. Smithson. Contr. Knowledge, 3 (Art. 4). 150 pp., pis. 1-12. 1853. Nereis boreali-americana: . . . Pt. 2. — Rhodospermeae. Smithson. Contr. Knowledge, 5 (Art. 5). 258 pp., pis. 13-36. Hassall, a. H. 1845. A History of the British Freshwater Algae . . . Vol. 1. viii + 462 + [1] pp. Vol. 2. 24 pp., 103 pis. London: Taylor, Walton, and Maberly. (Printing of 1852 and 1857 seen.) Hauck, F. 1883-1885. Die Meeresalgen Deutschlands und Oesterreichs. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz. 2nd ed. Bd. 2. xxiii + [1] + 575 pp., 236 figs., 5 pis. Leipzig: E. Kummer. Hedwig, J. 1798. Theoria generationis et fructificationis plantarum cryptogamicarum Linnaei retractata et aucta. x + 268 pp., pis. A, 1-42. Leipzig: Breitkopf. Heering, W. 1906. Die Siisswasseralgen Schleswig-Holsteins ... 1. Teil: Einleitung. — Hetero- kontae. Mitt. Bot. Staatsinst. Hamburg, 1905:61-150, 43 figs. 1914. Chlorophyceae. III. Ulothricales, Microsporales, Oedogonales. In A. Pascher, Die Siisswasser-flora Deutschlands, osterreichs und der Schweiz. Heft 6. iv + 250 pp., 384 figs. Jena: G. Fischer. 1921. Chlorophyceae IV. Siphonocladiales, Siphonales. In A. Pascher, Die Siisswasser- Flora Deutschlands, osterreichs und der Schweiz. Heft 7. iv + 103 pp., 94 figs. Jena: G. Fischer. Hertwig, O. 1876. Beitrage zur Kenntniss der Bildung, Befruchtung und Theilung des thier- ischen Eies. Morph. Jahrb., 1:347-434, pis. 10-13. Hieronymus, G. 1905. Bemerkungen iiber Chlamydomyxa lahyrintJiuloides Archer und Chlamydomyxa montana Lankester. Hedwigia, 44:137-157. HiRN, K. E. 1900. Monographie und Iconographie der Oedogoniaceen. Acta Soc. Sci. Fennicae, 27(l):i-iv, 1-394, [1], 64 pis. PAPENFUSS: CLASSIFICATION OF THE ALGAE 203 HOFMEISTER, W. 1867. Die Leliie von der Pflanzeiizelle. In Handbuch der physiologischen Botanik. Vol. 1(1). xii + 404 pp., 58 figs. Leipzig: W. Engelmann. HOLLANDE, A. 1942. Etude cytologique et biologique de quelques flagell^s libres. Volvocales, Crypto- monadines, Eugleniens, Protomastigines. Arch. Zool. Experiment. G6n., 83:1-268, 47 figs., pis. 1-17. HOLLENBERG, G. J. 1940. New marine algae from southern California. I. Amer. Journ. Bot., 27:868-877, 17 figs. HOVASSE, R. 1932. Contribution a I'etude des silicoflagelles . . . Bull. Biol. France et Belgique, 66:447-501, 11 figs., pis. 6, 7, HUBER, J. 1892. Contributions a la connaissance des Chaetophorees epiphytes et endophytes et leur aflJnites Ann. Sci. Nat. Bot., ser. 7, 16:265-359, pis. 8-18. Huber-Pestalozzi, G. 1938. Das Phytoplankton des Susswassers . . . In A. Thienemann, Die Binnenge- wiisser . . . Bd. 16, Teil 1. [5] + 342 pp., 395 figs, in text and on pis. 1-66. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung. 1941. Die Phytoplankton des Siisswassers. Systematik and Biologic. In A. Thiene- mann, Die Binnengewasser . . . Bd. 16, Teil 2(1). [1] +365 pp., 443 figs. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung. 1950. Das Phytoplankton des Siisswassers . . . Teil 3, Cryptophyceen, Chloromonadi- nen, Peridineen. In A. Thienemann, Die Binnengewasser . . . Bd. 16, Teil 3. ix + 310 pp., 300 figs. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung. HUSTEDT, F. 1930. Bacillariophyta (Diatomeae). In A. Pascher, Die Stisswasser-Flora Mittel- europas. 2nd ed. Heft. 10. viii + 466 pp., 875 figs. Jena: G. Fischer. Huxley, T. H. 1858. Appendix A. Report on the examination of specimens of bottom. In J. Dayman, Deep-Sea Soundings in the North Atlantic Ocean between Ireland and New- foundland, Made in H.M.S. "Cyclops" . . . 1857. London: Great Britain and Ireland. — Admiralty. (Not seen.) Hy, F. 1913. Les Characees de France. Soc. Bot. France, M^m. 26. 47 pp., 3 pis. Hygen, G. 1934. tJber den Lebenszyklus und die Entwicklungsgeschichte der Phaeosporeen. Versuche an Nemacystis divaricatus (Ag.) Kuck. Nyt. Mag. Naturvidensk., 74:187-268, 11 figs., 16 pis. Isiiikawa, M. 1921. Cytological studies on Porphyra tenera Kjellm. I. Bot. Mag. Tokyo, 35:206-218, 14 figs., pi. 4. 1924. On the phylogeny of Rhodophyceae. Bot. Mag. Tokyo, 38:159-167. (In Japanese with an English summary on p. 130.) Iyengar, M.O.P. 1936. Characiosiplion, a new member of the Chlorophyceae. (Preliminary note.) Journ. Indian Bot. Soc, 15:313-318, 10 figs., pi. 23. 204 A CENTURY OF PROGRESS /N THE NATURAL SCIENCES Iyengar, M.O.P., and T. V. Desikachary 1946. Mastigocladopsis jogensis gen. et. sp. nov., a new member of the Stigonemata- ceae. Proc. Indian Acad. Sci., 24:55-59, 13 figs., pi. 1. Iyengar, M.O.P., and K. R. Ramanathan 1940. On the reproduction of Anadyomene stellata (Wulf.) Ag. (Preliminary note.) Journ. Indian Bot. Soc, 19:175-176. 1941. On the life-history and cytology of Microdictyon tenuius (Ag.) Decsne. (Pre- liminary note.) Journ. Indian Bot. Soc, 20:157-159, 9 figs., pi. 3. Iyengar, M.O.P., and R. Subrahmanyan 1942. On reduction division and auxosporeformation in Cyclotella Meneghiniana Kiitz. (Preliminary note.) Journ. Indian Bot. Soc, 21:231-237, 14 figs., pi. 10. 1944. On reduction division and auxosporeformation in Cyclotella Meneghiniana Kutz. Journ. Indian Bot. Soc, 23:125-152, 63 figs., pis. 8, 9. Jahn, T. L. 1946. The euglenoid flagellates. Quart. Rev. Biol., 21:246-274, 6 figs. 1951. Euglenophyta. In G. M. Smith, ed.. Manual of Phycology. Pp. 69-81, figs. 18-20. Waltham, Mass.: Chronica Botanica Co. Janczewski, E. 1876. Notes sur le developpment du cystocarpe dans les Florid^es. M^m. Soc. Natl. Sci. Nat. Cherbourg, 20:109-144, pis. 3-5. Jane, F. W. 1946. A revision of the genus Harpochytrium. Jour. Linn. Soc. London, Bot., 53: 28-40, 28 figs. JAO, C. C. 1941. Studies on the freshwater algae of China. IX. Coelodiscaceae, a new family of the Chlorophyceae. Sinensia, 12:291-298, pi. 1. 1947. Coelodiscus buUaUis, sp. nov., a second species of the Coelodiscaceae. Bot. Bull. Acad. Sinica, 1:255-256, 1 fig. Johnson, T. 1894. The systematic position of the Bangiaceae. Nuova Not., 5:636-647. JORDE, I. 1933. Untersuschungen liber den Lebenszyklus von JJrospora Aresch. und Codiolum A. Braun. Nyt. Mag. Naturvidenskab., 73:1-19, 5 figs., pi. 1. JULLER, E. 1937. Der Generations- und Phasenwechsel bei Stigeoclonium subspinosum. Archiv Protistenk., 89:55-93, 21 figs. JUSSIEU, A. L. DE 1789. Genera plantarum ... (24) + Ixxii + 498 -f- [1] pp. Paris: Herissant. Kamptner, E. 1928. tJber das System und die Phylogenie der Kalkflagellaten. Archiv Protistenk., 64:19-43, 1 diagram. Karsten, G. 1899. Die Diatomeen der Kieler Bucht. Wiss. Meeresuntersuch. Abt. Kiel, N.F., 4:17-205, 219 figs. 1912. tJber die Reduktionsteilung bei der Auxosporenbildung von Surirella saxonica. Zeitschr. Bot., 4:417-426, pi. 7. PAPENFUSS: CLASSIFICATION OF THE ALGAE 205 Kabsten, G. (Cont.) 1928. Abteilung Bacillariophyta (Diatomeae). In A. Engler, Die natiirlichen Pflan- zenfamilien . . . 2nd ed. Bd. 2. pp. 105-303, figs. 93-424. Leipzig: W. Engel- mann. Kaulfuss, G. F. 1825. Erfahrungen iiber das Keimen der Charen, nebst andern Beitragen zur Kenntniss dieser Pflanzengattung. [8] + 92 pp., 1 pi. Leipzig: Publ. by author? Kent, W. S. 1880-1881. A Manual of the Infusoria: . . . Vol. 1. x + 472 pp. London: David Bogue. KiRCIINER, 0. 1878. Algen. 7n P. Cohn, Kryptogamen-Flora von Schlesien. Bd. 2(1). iv + [3] + 284 pp. Breslau: J. U. Kern's Verlag. 1898. Schizophyceae (Myxophyceae Stizenberger; Phycochromophyceae Rabenhorst; Cyanophyceae Sachs). In A. Engler and K. Prantl, Die natiirlichen Pflanzen- familien . . . Teil I, Abt. la. Pp. 45-92, figs. 48-62. Leipzig: W. Engelmann. Kjellman, p. R. 1880. Rhodospermeae et Pucoideae. In Enumerator plantae scandinaviae. Lund. (Not seen, cited from Kjeilman, 1883.) 1883. The algae of the Arctic Sea. K. Svensk. Vet.-Akad. Handl., Vol. 20 (5). 350 + [1] pp., 5 tables, 31 pis. 1890. Handbok i skandinaviens hafsalgflora. [4] + 103 pp., 17 figs. Stockholm: 0. L. Lamm. 1891-1893. Phaeophyceae. In A. Engler and K. Prantl, Die natiirlichen Pflanzen- familien . . . Teil 1, Abt. 2. Pp. 176-290, figs. 129-188. Leipzig: W. Engelmann. 1893. Dictyotales. In A. Engler and K. Prantl, Die natiirlichen Pflanzenfamilien . . . Teil 1, Abt. 2. Pp. 291-297, figs. 189-191. Leipzig: W. Engelmann. Klebahn, H. 1888. Ueber die Zygosporen einiger Conjugaten. Ber. Deutsch. Bot. Ges. 6:160-166, pi. 7. 1891. Studien iiber Zygoten L Die Keimung von Closterium und Cosmarium. Jahrb. Wiss. Bot, 22:415-443, pis. 13, 14. 1892. Studien iiber Zygoten II. Die Befruchtung von Oedogonium Boscii. Jahrb. Wiss. Bot., 24:235-267, pi. 3. 1896. Beitrage zur Kenntniss der Auxosporenbildung. I. Rhopalodia gihba (Ehrenb.) O. Miiller. Jahrb. Wiss. Bot., 29:595-654, pi. 10. Klebs, G. 1883. Uber die Organisation einiger Flagellaten-Gruppen und ihre Beziehung zu Algen und Infusorien. Untersuch. Bot. Inst. Tiibingen, 1(2) : 233-362, pis. 2, 3. 1884. Ein kleiner Beitrag zur Kenntniss der Peridineen. Bot. Zeit., 42:721-733, 737-745, pi. 10. 1892. Flagellatenstudien. I, II. Zeitschr. Wiss. Zool., 55:265-445, pis. 13-18. 1912. tJber Plagellaten- und Algen-ahnliche Peridineen. Verh. Naturhist.-Medizin. Ver. Heidelberg, N. P., 11:369-451, 15 figs., pi. 10. Knebel, G. 1935. Monographie der Algenreihe der Prasiolales, insbesondere von Prasiola crispa. Hedwigia, 75:1-120, 32 figs., 14 tables, 3 maps. Kniep, H. 1928. Die Sexualitat der niederen Pflanzen . . . vi + 544 pp., 221 figs., 23 tables. Jena: G. Pischer. 206 A CENTURY Of PROGRESS IN THE NATURAL SCIENCES Knight, Mabgery 1923. Studies in the Ectocarpaceae. I. The life-history and cytology of Pylaiella litoralis Kjellm. Trans. Roy. Soc. Edinburgh, 53(17) : 343-360, 6 pis. 1929. Studies in the Ectocarpaceae. II. The life-history and cytology of Ectocarpus siliculosus, Dillw. Trans. Roy. Soc. Edinburgh, 56(15): 307-332, 3 figs., 6 pis. 1947. John Lloyd Williams [Obituary]. Proc. Linn. Soc. London, 158:72-74. Koch, W. J. 1951. A study of the motile cells of Vaucheria. Journ. Elisha Mitchell Sci. Soc, 67: 123-131, pis. 3-5. KoFoiD, C. A., and T. Skogsbeeg 1928. The Dinoflagellata: the Dinophysoidae. In Reports on the Scientific Results of the Expedition to the Eastern Tropical Pacific, in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer "Albatross," . . . xxxv. Mem. Mus. Comp. Zool., Harvard College, Vol. 51. 766 pp., 103 figs., 31 pis. KoFoiD, C. A., and Olive Swezy 1921. The Free-Living Unarmored Dinoflagellata. Mem. Univ. Calif., Vol. 5. viii + 562 pp., 388 figs., 12 pis. KOLBE, R. W. 1948. Elektronenmikroskopische Untersuchungen von Diatomeenmembranen. Arkiv Bot. Vol. 33(17). 21 pp.. 5 figs., 10 pis. KoLKWiTZ, R., and H. Keiegee 1941-1944. Zygnemales. I7i L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2nd ed. Bd. 13, Abt. 2 (issued by R. Kolkwitz). [1] + 499 pp., figs. A-Z, 1-779, 4-9. Leipzig: Akad. Verlagsges. KORNMANN, P. 1938. Zur Entwicklungsgeschichte von Derdesia und Halicystis. Planta, 28:464-470, 4 figs. 1953. Der Formenkreis von Aciiietospora crinita (Carm.) nov. comb. Helgolander Wiss. Meeresuntersuch., 4:205-224, 14 figs. KORSCHIKOV, A. A. 1923. tJber zwei neue Organismen aus der Gruppe der Volvocales. Russ. Arkhiv ProtistoL, 2:177-178, pi. 7. 1926. On some new organisms from the groups Volvocales and Protococcales, and on the genetic relations of these groups. Archiv Protistenk., 55:439-503, 14 figs., pis. 4-12. 1929. Studies on the chrysomonads. I. Archiv Protistenk., 67:253-290, pis. 8-11. 1930. On the origin of the diatoms. Beih. Bot. Centr. 46 (Abt. 1, Heft 3) : 460-469, 1 fig. 1938. Volvocineae. (Not seen, cited from H. Skuja, 1939b.) KOSSINSKAJA, K. K. 1926. De novo cyanophycearum genere Tildenia mihi ad novam familiam Tildenia- ceae mihi ducto. Not. Syst. Inst. Crypt. Horti Bot. Principalis U.S.S.R., 4:76-88, pi. 2. (In Russian with a Latin resume.) Keiegee, W. 1933-1937. Die Desmidiaceen Europas mit Beriicksichtigung der aussereuropaischen Arten. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2nd ed. Bd. 13, Abt. 1 (issued by R. Kolkwitz). vi + 712 pp., 33 figs., 96 pis. Leipzig: Akad. Verlagsges. 1939. Die Desmidiaceen Europas mit Beriicksichtigung der aussereuropaischen Arten. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich PAPENFUSS: CLASSIFICATION OF THE ALGAE 207 Krieger, W. (Cont.) und der Schweiz. 2nd ed. Bd. 13, Abt. 1, Teil 2 (issued by R. Kolkwitz). 117 pp., 1 fig., pis. 97-142. Leipzig: Akad. Veiiagsges. KUCKUCK, P. 1891. Beitrage zur Kenntniss der Ectocarpus-Arter der Kieler Fohrde. Bot. Centr., 48:1-6, 33-41, 65-71, 97-104, 129-141, 6 figs. 1929. Fragmente einer Monographie der Phaeosporeen. Wiss. Meeresuntersuch., Abt. Helgoland, N. F., 17(4). 93 pp., 155 figs. (W. Nienberg, ed.) KUNIEDA, H. 1934. On the life-history ot Monostroma. Proc. Imp. Acad. Tokyo, 10:103-106, 12 figs. KuRssANOv, L. J., and N. M. Schemakhanova 1927. Sur la succession des phases nucleaires chez les algues vertes. Le cycle de developpement du Chlorochytrium Lemnae Cohn. Russ. Arkhiv Protistol., 6:131-146, pis. 9, 10. KiJTZiNG, F. T. 1833a. Beitrag zur Kenntniss fiber die Entstehung und Metamorphose der niedern vegetabilischen Organismen, nebst einer systematischen Zusammenstellung der hierher gehorigen niedern Algenformen. Linnaea, 8:335-384, pis. 6-8. 1833b. Synopsis diatomearum oder Versuch einer systematischen Zusammenstellung der Diatomeen. Linnaea, 8:529-620, pis. 13-19. 1843. Phycologia generalis . . . xvi + 1-144 + xvii-xxxii + 145-458 + [1] pp., 80 pis. Leipzig: Brockhaus. 1844. Die kieselschaligen Bacillarien oder Diatomeen. [1]+ 152 pp., 30 pis. Nord- hausen: Ferd. Forstemann. (Printing of 1865 seen.) 1845. Phycologia germanica . ., . x + [1] + 340 pp. Nordhausen: Publ. by author. 1849. Species algarum vi + 922 pp. Leipzig: Brockhaus. Kylin, H. 1916. tJber die Befruchtung und Reduktionsteilung bei Nemalion multifidum. Ber. Deutsch. Bot. Ges., 34(4) : 257-271, 7 figs. 1917. iJber die Entwicklungsgeschichte und die systematische Stellung der Tilopteri- deen. Ber. Deutsch. Bot. Ges., 35:298-310. 1918. Studien liber die Entwicklungsgeschichte der Phaeophyceen. Svensk Bot. Tidskr., 12:1-64, 30 figs. 1923. Studien liber die Entwicklungsgeschichte der Florideen. K. Svensk. Vet.-Akad. Handl., 63(11). 189 pp., 82 figs. 1925. The marine red algae in the vicinity of the Biological Station at Friday Harbor, Wash. Lunds Univ. Arsskr., N. F., Avd. 2, 21(9). 87 pp., 47 figs. 1928. Entwicklungsgeschichtliche Florideenstudien. Lunds Univ. Arsskr., N. F. Avd. 2, 24(4). 127 pp., 64 figs. 1930a. iJber die Entwicklungsgeschichte der Florideen. Lunds Univ. Arsskr., N. F. Avd. 2, 26(6). 104 pp., 56 figs. 1930b. Some physiological remarks on the relationship of the Bangiales. Bot. Not. 1930:417-420. 1932. Die Florideenordnung Gigartinales. Lunds Univ. Arsskr., N. F., Avd. 2, 28(8) 88 pp., 22 figs., 28 pis. 1933. tJber die Entwicklungsgeschichte der Phaeophyceen. Lunds Univ. Arsskr. N. F., Avd. 2, 29(7). 102 pp., 35 figs., 2 pis. 1935. uber einige kalkbohrende Chlorophyceen. K. Fysiogr. Sallskap. Lund. For handl., 5(19). 19 pp., 7 figs. 1937a. Anatomic der Rhodophyceen. In K. Linsbauer, Handbuch der Pflanzenanat omie. Bd. 6, Teilband 2: Algen (B, g). viii -|- 347 pp., 252 figs. Berlin: Gebr Borntraeger. 208 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES KXLIN, H. (Cont.) 1937b. Uber eine marine Porphyridium-Art. K. Fysiogr. Sallskap. Lund. Forhandl., 7(10). 5 pp., 1 fig. 1937c. Bemerkungen uber die Entwicklungsgeschichte einiger Phaeophyceen. Lunds Univ. Arsskr., N. F., Avd. 2, 33(1). 34 pp., 5 figs. 1938. Beziehungen zwischen Generationswechsel und Phylogenie. Archiv Protistenk., 90:432-447. 1940a. Die Phaeophyceenordnung Chordariales. Lunds Univ. Arsskr., N. F., Avd. 2, 36(9): 67 pp., 30 figs., 8 pis. 1940b. Zur Entwicklungsgeschicbte der Ordnung Fucales. Svensk Bot. Tidskr., 34: 301-314, 1 fig. 1943a. Zur Biochemie der Cyanophyceen. K. Fysiogr. Sallskap. Lund Forhandl., 13(7). 14 pp. 1943b. Verwandtschaftliche Beziehungen zwischen den Cyanophyceen und den Rhodo- phyceen. K. Fysiogr. Sallskap. Lund Forhandl., 13(17). 7 pp. 1944a. Die Rhodophyceen der schwedischen Westkiiste. Lunds Univ. Arsskr., N. F., Avd. 2, 40(2). 104 pp., 53 figs., 32 pis. 1944. Uber die Auxiliarzellen der Florideen. K. Fysiogr. Sallskap. Lund Forhandl., 14(20). 11 pp., 4 figs. 1947. Die Phaeophyceen der schwedischen Westkiiste. Lunds Univ. Arsskr., N. F., Avd. 2, 43(4). 99 pp., 61 figs., 18 pis. Lackey, J. B. 1939. Notes on plankton flagellates from the Scioto River (with description of new forms). Lloydia, 2:128-143, 38 figs. Lamarck, J. B. de, and A. P. de Cakdolle 1805. Flore frangaise . . . 3rd ed. Vol. 2 xli + 600 pp., map. Paris: Desray. Lamouhoux, J. V. F. 1805. Dissertations sur plusieurs especes de Fucus, peu connues on nouvelles, avec leur description en Latin et en Frangais. xxiv + 83 + [1] pp., 36 pis. Agen: Noubel. 1813. Essai sur les genres de la famille des thalassiophytes non articulees. Ann. Mus. Hist. Nat. [Paris], 20:21-47, 115-139, 267-293, pis. 7-13. Le Jolis, a. 1863. Liste des algues marines de Cherbourg. 168 pp., 6 pis. Paris: Bailliere et Fils. (1880 printing seen.) (Also publ. in Mem. Soc. Imp. Sci. Nat. Cherbourg 10, 1864.) Lemmermann, E. 1899. Planktonalgen. In H. Schauinsland, Ergebnisse einer Reise nach dem Pacific. Abhl. Nat. Ver. Bremen, 16(2) :313-398, pis. 1-3. 1901a. Beitrage zur Kenntniss der Planktonalgen. Ber. Deutsch. Bot. Ges. 19:85-95, pi. 4. 1901b. Silicoflagellatae. Ergebnisse einer Riese nach dem Pacific.-H. Schauinsland 1896/97. Ber. Deutsch. Bot. Ges., 19:247-271, pis. 10, 11. 1907-1910. Algen I (Schizophyceen, Flagellaten, Peridineen). In Kryptogamenflora der Mark Brandenburg und angrenzender Gebiete herausgegeben von dem Botanischen Verein der Provinz Brandenburg. Bd. 3. x + (l)-(29) + 712 pp., 816 figs. Leipzig: Gebr. Borntraeger. 1908. Flagellatae, Chlorophyceae, Coccosphaerales und Silicoflagellatae. In K. Brandt and C. Apstein, Nordisches Plankton, Botanischer Teil, 21. 40 pp., 135 figs. Kiel and Leipzig: Lipsius and Tischer. PAPENFUSS: CLASSIFICATION OF THE ALGAE 209 Lemmekmann, E. (Cont.) 1913. Eugleninae. In A. Pascher, Die Siisswasser-Flora Deutschlands, osterrelchs und der Schweiz. Heft. 2: Flagellatae II. Pp. 115-174, figs. 181-378. Jena: G. Fischer. 1914. Pantostomatinae, Protomastiginae, Distomatinae. In A. Pascher, Die Suss- wasser-Flora Deutschlands, osterreichs und der Schweiz. Heft 1: Flagellatae I. Pp. 28-138. Jena: G. Fischer. 1915. Tetrasporales. In A. Pascher, Die Siisswasser-Flora Deutschlands, osterreichs und der Schweiz. Heft. 5: Chlorophyceae II. Pp. 21-51, 33 figs. Jena: G. Fischer. Levbing, T. 1941. Die Meeresalgen der Juan Fernandez-Inseln. In C. Skottsberg, The Natural History of Juan Fernandez and Easter Island. Vol. 2. Pp. 601-670, 30 figs., pis. 49-53. Lewis, I. F. 1909. The life history of Griffithsia Bornetiana. Ann. Bot., 23:639-690, 2 figs., pis, 49-53. 1912. Alternation of generations in certain Florideae. Bot. Gaz., 53:236-242. Lindley, J. 1836. A Natural System of Botany; ...2nd ed. xxvi + 526 pp. London: Longman, et al. LiNDEMANN, E. 1928. Peridineae (Dinoflagellatae). In A. Engler and K. Prantl, Die naturlichen Pflanzenfamilien . . . 2nd ed. (issued by A. Engler) Bd. 2 (E. Jahn, ed.). Pp. 3-104, figs. 1-92. Leipzig: W. Engelmann. LiNDSTEDT, A. 1943. Die Flora der marinen Cyanophyceen der schwedischen Westkiite. 121 + [1] pp., 2 figs., 3 tables, 11 pis. Thesis. Lund: Publ. by author. Linnaeus, C. 1753. Species plantarum . . . Vol. 2. Pp. 561-1200+ [31]. Stockholm: Salvius. 1788. Systema naturae . . . 13th ed. (J. F. Gmelin, ed.). Vol. 1(6). Pp. 3021-3909. Leipzig: G. E. Beer. Lohmann, H. 1902. Die Coccolithophoridae, eine Monographie der Coccolithen bildenden Flagel- laten, zugleich ein Beitrag zur Kenntnis des Mittelmeerauftriebs. Archiv Protistenk., 1:89-165, pis. 4-6. 1904. Eier und sogenannte Cysten der Plankton-Expedition. Ergebn. Plankton Exped. Humboldt-Stiftung. Vol. 4(N). 61+ [1] pp., 7 pis. Kiel and Leipzig: Lipsius and Tischer. Longest, Pauline M. 1946. Structure of the cilia in Ectocarpus Mitchellae and Codium decorticatum. Journ. Elisha Mitchell Sci. Soc, 62:249-252, 3 figs. Luerssen, C. 1879. Handbuch der systematischen Botanik . . . Bd. 1. Kryptogamen. xii + 657 pp., 181 figs. Leipzig: Haessel. Luther, A. 1899. Ueber Chlorosaccus eine neue Gattung der Siisswasseralgen, nebst einigen Bemerkungen zur Systematik verwandter Algen. Bihang K. Svensk. Vet.- Akad. Handl., 24, Afd. 3, No. 13. 22 pp., 1 folding pi. Lyngbye, H. C. 1819. Tentamen hydrophytologiae danicae . . . xxxii + 248 pp., 70 pis. Copenhagen: Schultz. 210 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Macdonald, J. D. 1869. On the structure of the diatomaceous frustule, and its genetic cycle. Ann. Mag. Nat. Hist., ser. 4, 3:1-8, pi. 3. Mack, B. 1951. Morphologische und entwicklungsgeschichtliche Untersuchungen an Chryso- phyceen. oster. Bot. Zeitschr., 98:249-279, 3 figs. Mangenot, G. 1948. Sur les caracteres et la position systematique du genre Phyllo siphon. Compt. Rend. Acad. Sci. [Paris], 226:1103-1105. Manton, Irene, and B. Clakke 1951a. Electron microscope observations on the zoospores of Pylaiella and Laminaria. Journ. Exp. Bot., 2:242-246, 8 pis. 1951b. An electron microscope study of the spermatozoid of Fucus serratus. Ann. Bot., n. s., 15:461-471, pis. 20-24. Manton, Irene, B. Clarke, and A. D. Greenwood 1953. Further observations with the electron microscope on spermatozoids in the brown algae. Journ. Exp. Bot, 4:319-329, 2 figs., pis. 1-9. Madler, K. 1952. Charophyten aus dem Nordwestdeutschen Kimmeridge. Geol. Jahrb., 67:1-45, inc. pis. A, B, and 8 figs. 1953. Ein neues System der fossilen Charophyten. Flora, 140:474-484, 17 figs., pis. 2-5. Marc hand, L. 1895. Synopsis et tableau synoptique des families qui composent la classe des Phyco- phytes (Algues, Diatomees et Bacteriens). In Sous-regue des Cryptogames, 29 Embranchement. 20 pp., 1 chart. Paris: Soc. d'Edit. Sci. Martin, Margaret T. 1939. The structure and reproduction of CJiaetangium saccatuvi (Lamour.) J. Ag.-II. Female plants. Journ. Linn. Soc. Bot., 52:115-144, 38 figs., pi. 1. Mereschkowsky, C. 1879. Studien liber Protozoen des nordlichen Russland. Archiv Mikr. Anat., 16:153- 248, pis. 10, 11. Meyen, J. 1827. Ueber die Priestleysche griine Materie, wie iiber die Metamorphose des Proto- coccus viridis in Priestleya botryoides und in Ulva terrestris. Linnaea, 2: 388-409, pi. 7. Meyer, K. 1929. tJber die Auxosporenbildung bei Gomphonema geminatum. Archiv Protistenk., 66:421-435, pis. 15, 16. Migtjla, W. 1890-1897. Die Characeen. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz. 2nd edit. Bd. 5. xv + 765 pp., 149 figs. Leipzig: E. Kummer. (For dates of parts see Groves and Bullock-Webster, 1924, p. m.) 1895. Schizomycetes. In A. Engler and K. Prantl, Die naturlichen Pfiauzenfamilien . . . Teil 1. Abt. la. 44 pp., 47 figs. Leipzig: W. Engelmann. 1925. Charophyta-(Charales). In A. Pascher, Die Siisswasser-Flora Deutschlands, osterreichs und der Schweiz. Heft 11. Pp. 207-243, 14 figs. Jena: G. Fischer. PAPENFUSS: CLASSIFICATION OF THE ALGAE 211 Mitchell, Margaret O., and Frances G. Whitting 1892. On Splachnidium rugosum. Grev., the type of a new order of algae. In G. Murray, ed., Phycological Memoirs, Pt. 1. Pp. 1-10, pis. 1-3. London: Dulau and Co. MoBius, M. 1937. Geschichte der Botanik von den ersten Anfangen bis zur Gegenwart. vi + 458 pp. Jena: G. Fischer. MoEWUS, F. 1940. tJber die Sexualitat von Botrydium granulatum. Biol. Zentralbl., 60:484-498, 2 figs., 6 tables. MiJLLER, O. 1889. Durchbrechung der Zellwand in ihren Beziehungen zur Ortsbewegung der Bacillariaceen. Ber. Deutsch. Bot. Ges., 7:169-180, pi. 7. 1895. Ueber Achsen, Orientirungs- und Symmetrie-ebenen bei den Bacillariaceen. Ber. Deutsch. Bot. Ges., 13:222-234, pi. 20. MiJLLER, 0. F. 1773. Vermium terrestrium et fluviatilium, seu animalium infusoriorum, helminthi- corum et testaceorum . . . Vol. 1, pt. 1. [32] + 135 pp. Copenhagen and Leip- zig: Heineck and Faber. 1782. Icones plantarum sponte nascentium in regnis daniae et norvegiae . . . Vol. 5, fasc. 15. Pis. 841-900. Copenhagen: Moller. 1783. Strand-parlebandet och armbandet, tvanne microscopiska strandvaxter. K, Sv. Vet.-Akad. Handl., ser. 2, 4:80-85, pi. 3. 1786. Animalcula infusoria fluviatilia et marina . . . Opus hoc posthumum . . . cura 0. Fabricii. [8] + Ivi + 367 pp., 50 pis. Copenhagen: Moller. Murray, G. 1895. An Introduction to the Study of Seaweeds, xvi + 271 pp., 88 figs., 8 pis. London: Macmillan and Co. 1897. On the reproduction of some marine diatoms. Proc. Roy. Soc. Edinburgh, 21: 207-219, pis. 1-3. MtJRRAY, G., and V. H. Blackman 1898. On the nature of the coccospheres and rhabdospheres. Philos. Trans. Roy. Soc. London, 190 (ser. B): 427-441, pis. 15, 16. Murray, J. 1891. Deep-sea deposits. Rept. Sci. Results Voyage H.M.S. "Challenger" 1873-76. Pp. 257-258, figs. 19-21. Edinburgh: Neill and Co. Nageli, C. 1847. Die neuern Algensysteme und Versuch zur Begriindung eines eigenen Systems der Algen und Florideen. Neue Denkschr. Allgemein. Schweiz. Ges. Gesammt. Naturwiss., 9 (unnumbered art. No. 2). 275 pp., 10 pis. 1849. Gattungen einzelliger Algen physiologisch und systematisch bearbeitet. Neue Denkschr. Allgemein. Schweiz. Ges. Gesammt. Naturwiss., 10 (Art. 7). viii + 139 pp., 8 pis. 1861. Beitrage zur Morphologie und Systematik der Ceramiaceae. Sitzungsber. K. Bayer. Akad. Wiss. Miinchen, 2:297-415, 1 pi. Nasr, a. H. 1947. Synopsis of the marine algae of the Egyptian Red Sea coast. Bull. Faculty Sci. Fouad I Univ. no. 26. 155 pp., 25 figs., 14 pis. 212 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Newton, Linda M. 1953. Marine algae. In The John Murray Expedition 1933-34 Scientific Reports. Vol. 9, No. 5. Pp. 395-420, pis. 1-4. London: British Museum (Natural History). NiTZSCH, C. L. 1817. Beitrag zur Infusorienkunde oder Naturbeschreibung der Zerkarien und Bazillarien. viii + 128 pp., 6 pis. Halle: J. C. Hendel. Oehlkers, F. 1916. Beitrag zur Kenntnis der Kernteilungen bei den Charazeen. Ber. Deutsch. Bot. Ges., 34(4) : 223-227, 1 fig. Oltmanns, F. 1889. Beitrage zur Kenntniss der Fucaeeen. Bib). Bot. Vol. 3(14); 100 pp., 15 pis. 1898. Zur Entwickelungsgeschichte der Florideen. Bot. Zeit., 56:99-140, 2 figs., pis. 4-7. 1899. Ueber die Sexualitat der Ectocarpeen. Flora, 86:86-99, 16 figs. 1904. Morphologie und Biologic der Algen. Bd. 1. vi + 733 pp., 467 figs. Jena: G. Fischer. 1922a. Morphologie und Biologic der Algen. 2nd ed. Bd. 1. Chrysophyceae — Chloro- phyccae. vi + 459 pp., 287 figs. Jena: G. Fischer. 1922b. Morphologie und Biologic der Algen. 2nd ed. Bd. 2. Phaeophyceae — Rhodo- phyceac. iv + 439 pp., figs. 288-612. Jena: G. Fischer. OSTERHOUT, W. J. V. 1900. Befruchtung bei BatracTiospermum. Flora, 87:109-115, pi. 5. Palla, E. 1894. Ueber cine neuc, pyrenoidlose Art und Gattung der Conjugaten. Ber. Deutsch. Ges., 12:228-236, pi. 18. Papenfuss, G. F. 1933. Notes on the life-cycle of Ectocarpus siliculosus Dillw. Science, 77:390-391. 1935. Alternation of generations in Ectocarpus siliculosus. Bot. Gaz., 96:421-446, 13 figs., pis. 6, 7. 1946. Proposed names for the phyla of algae. Bull. Torrey Club, 73:217-218. 1947. Extension of the brown algal order Dictyosiphonales to include the Punctari- ales. Bull. Torrey Club, 74:398-402. 1950a. Review of the genera of algae described by Stackhousc. Hydrobiologia, 2: 181-208. 1950b. Culturing of marine algae in relation to problems in morphology. In J. Brunei, G. W. Prescott, and L. H. Tiffany, eds.. The Culturing of Algae, a Symposium. Pp. 77-95, 10 figs. The Charles F. Kettering Foundation. 1951a. Phaeophyta. In G. M. Smith, ed.. Manual of Phycology ... Pp. 119-158, figs. 29-37. Waltham. Mass.: Chronica Botanica Co. 1951b. Problems in the classification of the marine algae. Svensk Bot. Tidskr., 45: 4-11, 2 figs. Pascher, a. 1910. Chrysomonaden aus dem Hirschberger Grosstciche. Monogr. Abh. Intern. Rev. Gesammt. Hydrobiol. Hydrogr. Vol. 1. 66 pp., 3 pis. 1911a. Cyrto2)hora, eine neue tentakcltragende Chrysomonadc aus Franzensbad und ihre Verwandten. Ber. Deutsch. Bot. Ges., 29:112-125, 3 figs., pi. 6. 1911b. tJber die Beziehungen der Cryptomonaden zu den Algen. Ber. Deutsch. Bot, Ges., 29:193-203. PAPENFUSS: CLASSIFICATION OF THE ALGAE 213 Paschee, a. (Cont.) 1912a. t)ber Rhizopoden- und Palmellastadien bei Flagellaten (Chrysomonaden), nebst einer Ubersicht iiber die braunen Flagellaten. Archiv Protistenk., 25: 153-200, 7 figs., 1 chart, pi. 9. 1912b. Zur Gliederung der Heterokonten. (Kleine Beitrage zur Kenntnis unserer Mikroflora 3.) Hedwigla, 53:6-22, 8 figs. 1913a. Chrysomonadinae. In Pascher, Die Siisswasser-Flora Deutschlands, oster- reichs und der Schweiz. Heft 2: Flagellatae II. Pp. 7-95, figs. 1-150. Jena: G. Fischer. 1913b. Cryptomonadinae. I7i Pascher, Die Siisswasser-Flora Deutschlands, oster- reichs und der Schweiz. Heft 2: Flagellatae II. Pp. 96-114, figs. 151-180. Jena: G. Fischer. 1913c. Chloromonadinae. In Pascher, Die Siisswasserflora Deutschlands, osterreichs und der Schweiz. Heft 2: Flagellatae II. Pp. 175-181, figs. 379-387. Jena: G. Fischer. 1914. tJber Flagellaten und Algen. Ber. Deutsch. Bot. Ges., 32:136-160. 1915. Die Susswasser-Flora Deutschlands, osterreichs und der Schweiz. Heft 5: Chlorophyceae II, Einleitung. Pp. 1-20. Jena: G. Fischer. 1916a. Zur Auffassung der farblosen Flagellatenreihen. Ber. Deutsch. Bot. Ges., 34(7):440-447. 1916b. tJber eine neue Amobe — Dinamoehe (varians) — mit dinoflagellatenartigen Schwarmern. Archiv Protistenk., 36:118-136, 4 figs., pi. 10. 1917. Flagellaten und Rhizopoden in ihren gegenseitigen Beziehungen. Versuch einer Ableitung der Rhizopoden. Archiv Protistenk., 38:1-88, 65 figs. 1921. tJber die tJbereinstlmmung zwischen den Diatomeen, Heterokonten und Chryso- monaden. Ber. Deutsch. Bot. Ges., 39:236-248, 6 figs. 1924. Zur Homologisierung der Chrysomonadencysten mit den Endosporen der Dia- tomeen. Archiv Protistenk., 48:196-203, 4 figs. 1925a. Heterokontae. In Pascher, Die Siisswasser-Flora Deutschlands, osterreichs und der Schweiz. Heft 2. iv + 118 pp., 96 figs. Jena: G. Fischer, 1925b. Die braune Algenreihe der Chrysophyceen. Archiv Protistenk., 52:489-564, 56 figs., pi. 15. 1927a. Die braune Algenreihe aus der Verwandtschaft der Dinoflagellaten (Dinophy- ceen). Archiv Protistenk., 58:1-54, 38 figs. 1927b. Die Siisswasserflora Deutschlands, osterreichs und der Schweiz. Heft 4. 506 pp., 451 figs. Jena: G. Fischer. 1929a. Beitrage zur allgemeinen Zellehre. I. Doppelzellige Flagellaten und Parallel- entwicklungen zwischen Flagellaten und Algenschwarmern. Archiv Protis- tenk., 68:261-304, 21 figs. 1929b. tJber die Teilungsvorgange bei einer neuen Blaualge: Endonema. Jahrb. Wiss. Bot, 70:329-347, 10 figs. 1930a. Zur Kenntnis der heterokonten Algen. Archiv Protistenk., 69:401-451, 45 figs., pi. 21. 1930b. Zur Verwandtschaft der Monadaceae mit den Chrysomonaden: eine gehause- bewohnende, farblose Chrysomonade. Ann. Protistol., 2(4): 157-168, 6 figs. 1930c. Ober einen griinen, assimilationsfahigen plasmodialen Organismus in den Blat- tern von Sphagnum. Archiv Protistenk., 72:311-358, 27 figs., pis. 20, 21. 1931. Systematische tlbersicht iiber die mit Flagellaten in Zusammenhang steheden Algenreihen und Versuch einer Einreihung dieser Algenstamme in die Stamme des Pflanzenreiches. Beih. Bot. Centr., 48 (Abt. II): 317-332. 1932. tJber die Verbreitung endogener bzw. endoplasmatisch gebildeter Sporen bei den Algen. Beih. Bot. Centr., 49:293-308, 13 figs. 214 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Paschek, a. (Cont.) 1937-1939. Heterokonten. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, 5sterreich und der Schweiz. 2nd ed. Bd. 2 (issued by R. Kolkwitz). x + ii + 1092 pp., 912 figs. Leipzig: Akad. Verlagsges. 1939. tjber gelegentliche Synzoosporenbildungen bei Algen und Uber die Verbreitung synzoosporer Organisationen. Beih. Bot. Centr., 59 (Abt. A): 389-408, 20 figs. 1953. Lebenslauf und Arbeitenverzeichnis. Archiv Protistenk., 98:iii-xxxii. Peck, R. E. 1934. The North American trochiliscids, Paleozoic Charophyta. Journ. Paleont, 8:83-119, 2 figs., pis. 9-13. 1946. Fossil Charophyta. Amer. Mid. Nat., 36:275-278, 6 figs. Pebsidsky, B. M. 1929. The development of the auxospores in the group of the Centricae (Bacillaria- ceae). 11 pp., 22 figs., 1 double pi. Moskow. (Not seen, cited from the review by P. Schmidt in Zeitschr. Bot., 22[9] : 459-461, 1930.) 1935. The sexual process in Melosira varians. Beih. Bot. Centr., 53 (Abt. A) : 122-132, 23 figs. Perty, M. 1852. Zur Kenntniss kleinster Lebensformen nach Bau, Funktionen, Systematik, mit Specialverzeichniss der in der Schweiz beobachteten. viii + 228 pp., 17 pis. Bern: Jent and Reinert. Petersen, J. B. 1918. Om Synura Uvella Stein og nogle andre Chrysomonadiner. Videnskab. Medd. Dansk Naturhist. Foren. K30benhavn, 69:345-357, pi. 5. 1929. Beitrage zur Kenntnis der Flagellatengeisseln. Bot. Tidsskr. 40(5) :373-389, 11 figs. Pfitzer, E. 1869. Ueber Bau und Zelltheilung die Diatomaceen. Sitzungsber. Niederrhein. Ges. Natur. Heilk. Bonn, 26:86-89. 1871. Untersuchungen fiber Bau und Entwicklung der Bacillariaceen (Diatomaceen). In J. Hanstein, Botanische Abhandlungen aus dem Gebiet der Morphologic und Physiologie. Heft 2. vi + 189 pp., 6 pis. Bonn: A. Marcus. PlA, J. 1927. Thallophyta. In M. Hirmer, Handbuch der Paliiobotanik. Bd. 1. Pp. 31-136, figs. 14-129. Munich and Berlin: Oldenbourg. Pitelka, Dorothy R. 1949. Observations on flagellum structure in Flagellata. Univ. Calif. Publ. Zool., 53(11) :377-429, incl. pis. 37-46, 3 figs. Poche, F. 1913. Das System der Protozoa. Archiv Protistenk., 30:125-321, 1 fig. PococK, Mary A. 1954. Two multicellular motile green algae, Volvulina Playfair and Astrephomene, a new genus. Trans. Roy. See. So. Afr., 34:103-127, 30 figs., pi. 2. Pkescott, G. W. 1951. Algae of the Western Great Lakes Area Exclusive of Desmids and Diatoms, xiii + 946 pp., incl. 136 pis., text figs. 1-9, tables 1-7, 13 pp. (52-64) of unnum- bered illus. Cranbrook Inst. Sci. Bull., No. 31. Pringsheim, E. G. 1944. Some aspects of taxonomy in the Cryptophyceae. New Phytol., 43:143-150. 1948a. Taxonomic problems in the Euglenineae. Biol. Rev., 23:46-61, 4 figs. PAPENFUSS: CLASSIFICATION OF THE ALGAE 215 Pbingsheim, E. G. (Cont.) 1948b. The loss of chromatophores in Euglena gracilis. With a cytological contribution by R. Hovasse. New Phytol., 47:52-87, 13 figs. 1949. The relationship between bacteria and Myxophyceae. Bact. Rev., 13 : 47-98. Pbingsheim, E. G. and Olga 1952. Experimental elimination of chromatophores and eye-spot in Euglena gracilis. New Phytol., 51:65-76. Pbingsheim, E. G., and R. Hovasse 1950. Les relations de parents entre Astasiac^es et Euglenac6es. Arch. Zool. Exp^r. Gen., 86(8):499-549, 14 figs., pi. 16. Pbingsheim, N. 1855 iJber die Befruchtung der Algen. Ber. Verb. K. Preuss. Akad. Wiss. Berlin, 1855:133-165, 1 folding pi. 1856. uber die Befruchtung und den Generationswechsel der Algen. Monatsber. K. Preuss. Akad. Wiss. Berlin, 1856:225-237, 1 pi. 1860. Beitrage zur Morphologie und Systematik der Algen. III. Die Coleochaeteen. Jahrb. Wiss. Bot. 2:1-38, pis. 1-6. 1863a. tJber die Vorkeime der Charen. Monatsber. K. Preuss. Akad. Wiss. Berlin, 1862:225-231. 1863b. Ueber die Vorkeime und die nacktfiissigen Zweige der Charen. Jahrb. Wiss. Bot., 3:294-324, pis. 9-13. 1870. Uber Paarung von Schwarmsporen, die morphologische Grundform der Zeu- gung im Pflanzenreiche. Monatsber. K. Preuss. Akad. Wiss. Berlin, 1869: 721-738, 1 folding pi. Printz, H. 1927. Chlorophyceae. In A. Engler and K. Prantl, Die natiirlichen Pflanzenfamilien . . . 2nd ed. Vol. 3. iv + 463 pp., 366 figs. Leipzig: W. Engelmann. Rabenhobst, L. 1847. Deutschlands Kryptogamen-Flora . . . Bd. 2, Abt. 2: Algen. xix + [1] + 216 pp. Leipzig: E. Kummer. 1853. Die Siisswasser-Diatomaceen. (Bacillarien.) Fiir Freude der Mikroskopie. xii + 72 pp., 10 pis. Leipzig: E. Kummer. 1863. Kryptogamen-Flora von Sachsen, der Ober-Lausitz, Thiiringen und Nordbohmen . . . Abt. 1. XX + 653 pp., illustr. with text figs. Leipzig: E. Kummer. 1864. Flora europaea algarum aquae dulcis et submarinae. Sectio I. Algas dia- tomaceas complectens. 359 pp., illustr. with text figs. Leipzig: E. Kummer. 1865. Flora europaea algarum aquae dulcis et submarinae. Sectio II. Algas phyco- chromaceas complectens. 319 pp., illustr. with text figs. Leipzig: E. Kummer. 1868. Flora europaea algarum aquae dulcis et submarinae. Sectio III. Algas chloro- phyllophyceas, melanophyceas et rhodophyceas complectens. xx + 461 pp., illustr. with text figs. Leipzig: E. Kummer. Ralfs, J. 1848. The British Desmidieae. xxii + 226 pp., 35 pis. London: Reeve, Benham, and Reeve. Reinke, J. 1877. tJber das Wachsthum und die Fortpflanzung von Zanardinia collaris Crouan. (Z. prototypus Nardo.) Monatsber. K. Preuss. Akad. Wiss. Berlin, 1876: 565-578, 1 pi. 1878. Entwicklungsgeschiehtliche Untersuchungen iiber die Cutleriaceen des Golfs von Neapel. Nova Acta K. Leop.-Carol. Deutsch. Akad. Naturforsch., 40(2): 59-96, pis. 8-11. 216 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Rosenberg, Marie 1930. Die geschlechtliche Fortpflanzung von Botrydium granulatum Grev. osterr. Bot. Zeitschr., 79:289-296, 4 figs., 1 pi. Rosenberg, T. 1933. Studien iiber Rhodomelaceen und Dasyaceen. 87 pp., 25 figs. Thesis. Lund: Publ. by author. ROSENVINGE, L. K. 1909. The marine algae of Denmark . . . Part 1. Introduction. Rhodophyceae I. (Bangiales and Nemalionales.) K. Dansk. Vidensk. Selsk. Skrifter, 7. Raekke, Naturvidensk. Math. Afd., 7(1). 151 pp., 73 figs., 2 maps, 2 pis. ROSTAFINSKI, J. 1882. UHydrurus et ses affinit^s. Ann. Sci. Nat. Bot., ser. 6, 14:5-25, pi. 1. RosTAFiNSKi, J., and M. Woronin 1877. Ueber Botrydium granulatum. Bot. Zeit., 35:649-671, pis. 7-11. RUPRECHT, p. J. 1851. Tange des Ochotskischen Meeres. In A. Th. v. Middendorff's Sibirische Reise, Botanik. Bd. 1(2). Pp. 191-435, pis. 9-18. Leningrad: K. Akad. Wiss. 1855. tJber das System der Rhodophyceae. Mem. Acad. Imp. Sci. Saint-Petersbourg, ser. 6, Sci. Nat. Bot, 7:25-54, 1 pi. Sachs, J. 1874. Lehrbuch der Botanik ... 4th ed. xvi + 928 pp., 492 figs. Leipzig: W. Engelmann. 1875. Textbook of botany . . . (1st Eng. ed., trans, and annotated by A. W. Bennett and W. T. T. Dyer.) xii + 858 pp., 461 figs. Oxford: Clarendon Press. Sauvageau, C. 1896a. Sur la conjugaison des zoospores de VEctocarpus siliculosis. Compt. Rend. Acad. Sci. [Paris], 123:431-433. 1896b. Remarques sur la reproduction des Pheospor^es et en particulier des Ectocarpus. Ann. Sci. Nat. Bot, ser 8, 2:223-274. 1897. La copulation isogamique de \'Ectoca7-pus siliculosus est-elle apparente ou reele? Mem. Soc. Natl. Sci. Nat Math. Cherbourg, 30:293-304. 1915. Sur la sexualite heterogamique d'une Laminaire (Saccorhiza bulbosa). Compt. Rend. Acad. Sci. [Paris], 161:769-799, 3 figs. 1926. Sur un nouveau type d'alternance de generations chez les algues brunes; les Sporochnales. Compt Rend. Acad. Sci. [Paris], 182:361-364. 1931. Sur quelques algues pheosporees de la rade de Villefranche (Alpes-Maritimes). Bull. Stat Biol. Arcachon, 28:7-168, 32 figs. Schechner-Fries, Margarete 1934. Der Phasenwechsel von Valonia utricularis (Roth) Ag. osterr. Bot. Zeitschr., 83:241-254, 3 figs. Scherffel, a. 1901. Kleiner Beitrag zur Phylogenie einiger Gruppen niederer Organismen. Bot. Zeit, 59:143-158, pi. 6. 1911. Beitrag zur Kenntnis der Chrysomonadineen. Archiv Protistenk., 22:229-344, pi. 16. 1924. Tiber die Cyste von Monas. Archiv Protistenk., 48:187-195, 6 figs. 1927. Beitrag zur Kenntnis der Chrysomonadineen. II. Archiv Protistenk., 57:331- 361, 3 figs., pi. 15. Schiffner, v. 1926. Die Bxistenzgrunde der Zellbildung und Zellteilung, der Vererbung und Sex- aulitat . . , iv + [1] -f 160 pp. Jena: G. Fischer. PAPENFUSS: CLASSIFICATION OF THE ALGAE 217 SCHILLEB, J. 1925a. Die planktontischen Vegetationeii des adriatischen Meei-es. A. Die Coccolitho- phoriden-Vegetatiou in den Jahren 1911-14. Arcliiv Protistenk., 51:1-130. 24 figs., 11 tables, pis. 1-9. 1925b. Die planktontischen Vegetationen des adriatischen Meeres. B. Chrysomonadina, Heterokontae, Cryptomonadina, Eugleninae, Volvocales. 1. Systematischer Teil. Archiv Protistenk., 53:59-123, 30 figs., pis. 3-6. 1926. tJber Fortpflanzung, geissellose Gattungen und die Nomenklatur der Cocco- lithophoraceen nebst Mitteilung iiber Copulation bei Dinohryon. Archiv Protistenk., 53:326-342, 8 figs. 1930. Coccolithineae. In L. Rabenhorst's Kryptogamen-Flora von Deutschland, oster- reich und der Schweiz. 2nd ed. Bd. 10, Abt. 2 (issued by R. Kolkwitz). Pp. 89-273, figs. 1-139, A-F. Leipzig: Akad. Verlagsges. 1931-1933. Dinoflagellatae (Peridineae). 7?! L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2nd ed. Bd. 10, Abt. 3, Teil 1 (issued by R. Kolkwitz). vi + 617 pp., 631 figs. Leipzig: Akad. Verlagsges. 1935-1937. Dinoflagellatae (Peridineae). In L. Rabenhorst's Kryptogamen-Flora von Deutschland, osterreich und der Schweiz. 2ud ed. Bd. 10, Abt. 3, Teil 2 (issued by R. Kolkwitz). vii + 589 -f [IJ PP-, 612 figs. Leipzig: Akad. Verlagsges. SCIIILLIXG, A. J. 1913. Dinoflagellatae (Peridineae). In A. Pascher, Die Siisswasserflora Deutschlands, osterreichs und der Schweiz. Heft 3. iv + 66 pp., 69 flgs. Jena: G. Fischer. SCIIMIDLE, W. 1903. Bemerkungen zu einigen Siisswasseralgen. Ber. Deutsch. Bot. Ges., 21:346-355, pi. 18. Schmidt, 0. C. 1937a. Choristocarpaceen und Discosporangiaceen. Hedwigia, 77(1) :l-4. 1937b. Die Masonophyceen, eine neue Familie der Braunalgen. Hedwigia, 77(1) :5-6. 1938. Beitrage zur Systematik der Phaeophyten I. Hedwigia, 77(5, 6) : 213-230. SCHMITZ, C. J. F. 1879a. Ueber griine Algen im Golf von Athen. Sitzungsber. Naturf. Ges. Halle (Sitzung am. 30. Nov. 1878), pp. 17-23. (Reprinted under the title "Ueber grune Algen aus dem Golf von Athen" in Bot. Zeit, 37:167-175, 1879.) lS79b. Untersuchungen iiber die Fruchtbildung der Squamarieen. Sitzungsber. Nie- derrhein. Ges. Natur- u. Heilkunde Bonn (in Verb. Naturhist. Ver. Preuss. Rheinlande u. Westfalens), 36:376-377. 1879c. Untersuchungen iiber die Zellkerne der Thallophyten. Sitzungsber. Nieder- rhein. Ges. Natur- u. Heilkunde Bonn, 1878:345-376. 1882. Die Chromatophoren der Algen. Vergleichende Untersuchungen iiber Bau und Entwicklung der Chlorophyllkorper und analogen Farbstoffkorper der Algen. iv + 180 pp., 2 pis. Bonn: Max Cohen and Sohn. (Also in Verh. Naturhist. Ver. Preuss. Rheinl. Westfal., 40(1883) : 1-180, pis. 1, 2.) 1883. Untersuchungen liber die Befruchtung der Florideen. Sitzungsber. K. Preuss. Akad. Wiss. Berlin, 1883(1) :215-25S, pi. 5. 1889. Systematische tJbersicht der bisher bekannten Gattungen der Florideen. Flora, 72:435-456, pi. 21. ScHMiTZ, C. J. F., and P. Hauptfleisch 1896-1897. Rhodophyceae. In A. Engler and K. Prantl, Die natiirlichen Pflanzen- familien . . . Teil 1, Abt. 2. Pp. 298-544, figs. 192-288. Leipzig: W. Engelmann. SCHULZ, P. 1928. Beitrage zur Kenutnis fossiler und rezenter Silicoflagellaten. Bot. Archiv, 21:225-292, 83 figs. 218 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES SCHULZE, K. L. 1939. Cytologische Untersuchungen an Acetabularia mediterranea und Acetaiularia wettsteinii. Archiv Protistenk., 92:179-225, 20 figs., pis. 11, 12. ScHtrssNiG, B. 1925. Betrachtungen liber das System der niederen Pflanzen. Verh. Zool.-Bot. Ges. Wien, 74-75; 196-272. 1930. Der Generations- und Phasenwechsel bei den Clalorophyceen. (Ein historicher Rucliblick.) osterr. Bot. Zeitschr., 79:58-77. 1932. Der Generations- und Phasenwechsel bei den Chlorophyceen III. osterr. Bot. Zeitschr., 81:296-298. 1938. Der Kernphasenwechsel von Valonia utricularis (Roth) Ag. Planta., 28:43-59, 3 figs. 1939. Ein Beitrag zur Eutwiklungsgeschichte von Caulerpa proUfera. Bot. Not. 1939: 75-96, 6 figs. 1950. Die Gametogenese von Codium decorticatum (Woodw.) Howe. Svensk Bot. Tidskr., 44:55-71, 2 figs., pis. 1, 2. SCHUTT, F. 1888. Ueber die Diatomeengattung Chaetoceros. Bot. Zeit, 46:161-170, 177-184, pi. 3. 1895. Die Peridineen der Plankton-Expedition. I. Theil. Studien iiber die Zellen der Peridineen. Ergebnisse der Plankton-Expedition der Humboldt-Stiftung. Bd. iv. M. a. A. 170 pp., 27 pis. Kiel and Leipzig: Lipsius and Tischer. 1896. Gymnodiniaceae, Prorocentraceae, Peridiniaceae, Bacillariaceae. In A. Engler and K. Prantl, Die natiirlichen Pflanzenfamilien . . . (continued by A. Engler). Teil 1, Abt. lb. 153 pp., 282 figs. Leipzig: W. Engelmann. ScHWARz, Elisabeth 1932. Beitrage zur Entwicklungsgeschichte der Protophyten. IX. Der Formwechsel von Ochrosphaera neapoUtana. Archiv Protistenk., 77:434-462, 7 figs., pi. 13. Senn, G. 1900. Flagellata. In A. Engler and K. Prantl, Die natiirlichen Pflanzenfamilien . . . Teil I, Abt la. Pp. 93-188, figs. 63-140. Leipzig: W. Engelmann. Setchell, W. a. 1929. The genus Microdictyon. Univ. Calif. Publ. Bot., 14(20) : 453-588, 105 figs. Setchell, W. A., and N. L. Gardner 1919. The marine algae of the Pacific coast of North America. Pt. 1, Myxophyceae. Univ. Calif. Publ. Bot, 8(1) : 1-138, incl. pis. 1-8. 1920. The marine algae of the Pacific coast of North America. Pt. 2. Chlorophyceae. Univ. Calif. Publ. Bot, 8(2) :139-375, incl. pis. 9-33. 1922. Phycological contributions. VI. New species of Ectocarpus. Univ. Calif. Publ. Bot, 7(11): 403-427, incl. pis. 45-49. 1925. The marine algae of the Pacific coast of North America. Pt. 3. Melanophyceae. Univ. Calif. Publ. Bot, 8(3) :383-898, incl. pis. 34-107. Seybold, a., K. Egle, and W. Hijlsbruch 1941. Chlorophyll- und Carotinoidbestimmungen von Siisswasseralgen. Bot. Archiv, 42:239-253, 6 tables. SlEBOLD, C. T. VON 1848. Wirbellose Thiere. In C. T. Siebold and H. Stannius, Lehrbuch der vergleichen- den Anatomie. Vol. 1. xiv -f 679 + [1] pp. Berlin: Veit and Co. 1849. Ueber einzellige Pflanzen und Thiere. Zeitschr. Wiss. Zool., 1:270-294. PAPENFUSS: CLASSIFICATION OF THE ALGAE 219 SiLVA, P. C. 1952. A review of nomenclatural conservation in the algae from the point of view of the type method. Univ. Calif. Publ. Bot, 25(4) : 241-323. SiLVA, P. C, and R. C. Starr 1953. Difficulties in applying the international code of botanical nomenclature to certain unicellular algae, with special reference to Chlorococcum. Svensk Bot. Tidskr., 47:235-247. Singh, R. N. 1945. Nuclear phases and alternation of generations in Draparnaldiopsis indica Bharadwaja. New Phytol., 44:118-129, 49 figs. 1947. Fritschiella tuherosa lyeng. Ann. Bot., n. s., 11:159-164, 11 figs. Sjostedt, L. G. 1926. Floridean studies. Lunds Univ. Arsskr., N. F., Avd. 2, 22(4). 95 pp., 41 figs. Skottsberg, C. 1907. Zur Kenntnis der subantarktischen und antarktischen Meeresalgen. I. Phaeo- phyceen. Wiss. Ergebn. Schwed. Sudpolar-Exped. 1901-1903. Bd. 4(6). 172 pp., 187 figs., 10 pis., 1 map. Stockholm: Norstedt and Soner. Skuja, H. 1938. Die phylogenetischen Entwicklungsrichtungen bei den Protisten. Acta Biol. Latvica, 8:1-26. 1939a. Versuch einer systematischen Einteilung der Bangioideen oder Protoflorideen. Acta Hort. Bot. Univ. Latviensis, 11-12:23-40. 1939b. Beitrag zur Algenflora Lettlands II. Acta Hort. Bot. Univ. Latviensis, 11-12: 41-169, pis. 1-11. 1948. Taxonomie des Phytoplanktons einiger Seen in Uppland, Schweden. Symb. Bot. Upsal., Vol. 9(3). 399 pp., 39 pis. 1950. Korperbau und Reproduktion bei Dinodryon Borgei Lemm. Svensk Bot. Tidskr., 44:96-107, 1 fig. Smith, G. M. 1920. Phytoplankton of the Inland Lakes of Wisconsin. Pt. I, Myxophyceae, Phaeo- phyceae, Heterokonteae, and Chlorophyceae exclusive of the Desmidiaceae. Wis. Geol. Nat. Hist. Surv., Sci. Ser. No. 12. 243 pp., 2 tables, 51 pis. Madison, Wis.: Publ. by the State. 1930. Observations on some siphonaceous green algae of the Monterey Peninsula. In Contributions to Marine Biology. Pp. 222-233, 3 figs. Stanford Univ. Press. 1933. The Fresh-water Algae of the United States, xi + 716 pp., 449 figs., 2 tables. New York and London: McGraw-Hill. 1938. Cryptogamic Botany. Vol. I. viii -J- 545 pp., 299 figs. New York and London: McGraw-Hill. 1944. Marine Algae of the Monterey Peninsula California, vii + [1] + 622 pp., incl. 98 pis. Stanford Univ. Press. 1950. The Fresh-water Algae of the United States. 2nd ed. vii + 719 pp., 559 figs., 3 tables. New York, Toronto, London: McGraw-Hill. SOLMS-LAtTBACH, H. 1867. Ueber die Fruchtentwickelung von Batrachospermum. Bot. Zeit., 25:161-167, 169-171, pi. 4. Stackhouse, J. 1795-1801. Nereis britannica . . . xi + 112 -f [7] pp., 17 +A-G pis. Bath: Hazard. 1809. Tentamen marino-cryptogamicum . . . M6m. Soc. Imp. Nat. Moscou, 2:50-97. 1816. Nereis britannica . . . 2nd ed. xii + 68 pp., 20 pis. Oxford: Collingwood. 220 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Staxier, R. Y., and C. B. van Niel 1941. The main outlines of bacterial classification. Journ. Bact., 42:437-466, 2 figs. Stein, F. von 1878. Der Organismus der Infusionsthiere . . . Abt. 3. Die Naturgeschichte der Flagellaten oder Geisselinfusorien. I. Halfte. Den noch nicht abgeschlossenen allgemeinen Theil nebst Erklarung der sammtlichen Abbildungen enthal- tend. X + 154 pp., 24 pis. Leipzig: W. Engelmann. 1883. Der Organismus der Infusionstheire . . . Abt. 3. Die Naturgeschichte der Flagellaten oder Geisselinfusorien. II. Halfte. Die Naturgeschichte der ar- throdelen Flagellaten. 30 pp., 25 pis. Leipzig: W. Engelmann. Steinecke, F. 1932. Algologische Notizen II. Heteroclendron Pascheri, Euglenocapsa oehracea, Stylodinium cerasiforme. Archiv Protistenk, 76:589-594, 3 figs. Stizenbergeb, E. 1860. Dr. Ludwig Rabenhorst's Algen Sachsens resp. Mitteleuropa's Decade I-C. Systematisch geordnet (mit Zugrundelegung eines neuen Systems). 41 pp. Dresden: Heinrich. Stosch, H. a. 1951a. Entwicklungsgeschlichtliche Untersuchungen an zentrischen Diatomeen I. Die Auxosporenbildung von Melosira varians. Archiv. Mikrobiol., 16:101-135, inc. 2 pis. and 22 figs. 1951b. Zur Entwicklungsgeschichte zentrischer Meeresdiatomeen. Naturwiss., 38: 191-192. Strain, H. H. 1948. Occurrence and properties of chloroplast pigments. Carnegie Inst. Washington Year Book No. 47, pp. 97-100. 1951. The pigments of algae. In G. M. Smith, ed.. Manual of Phycology ... Pp. 243-262, 7 tables. Waltham, Mass.: Chronica Botanica Co. Strasbubger, E. 1887. Das botanische Practicum. 2nd ed. xxxvi + 685 pp., 193 figs. Jena: G. Fischer. 1897. Kernteilung und Befruchtung bei Fucus. Jahrb. Wiss. Bot, 30:351-374, pis. 17, 18. Strehlovt, K. 1929. tJber die Sexualitat einiger Volvocales. Zeitschr. Bot., 21:625-692, 17 figs. Stromfelt, H. F. G. 1886. Om algvegetationen vid Islands kuster. 89 pp., 1 table, 3 pis. Thesis (Upsala). Gothenburg: Publ. by author. Sturch, H. H. 1926. Choreocolax Polysiplioniae Reinsch. Ann. Bot., 40:585-605, 15 figs. Subbahmanyan, R. 1946. On the occurrence of microspores in some centric diatoms of the Madras coast. Journ. Indian Bot. Soc, 25:61-66, 12 figs., pi. 4. 1947. On somatic division, reduction division, auxospore-formation and sex-differen- tiation in Navicula haloj)Mla (Grun.) CI. M. O. P. Iyengar Comm. Vol., Journ. Indian Bot. Soc, pp. 239-266, 81 figs., pis. 1, 2. SUNESON, S. 1947. Notes on the life-history of Monostroma. Svensk Bot. Tidskr., 41:235-246, 2 figs., 1 table. PAPENFUSS: CLASSIFICATION OF THE ALGAE 221 SVEDELIUS, N. 1915. Zytologisch-entwicklungsgeschichtliche Studien iiber Scinaia furcellata ein Beitrag zur Frage der Reduktionsteilung der nicht tetrasporenbildenden Florideen. Nova Act Reg. Soc. Sci. Upsal., ser. 4, 4(4). 55 pp., 32 figs. 1916. Das Problem des Generationswechsels bei den Florideen. Naturwiss. Wochen- schr., N. F., 15:353-359, 372-379, 14 figs. 1942. Zytologisch-entwicklungsgeschichtliche Studien iiber Galaxaura eine diplo- biontische Nemalionales-Gattung. Nova Acta Reg. Soc. Sci. Upsal., ser. 4, 13(4). 154 pp., 80 figs. 1953. Critical studies on some species of Galaxaura from Hawaii. Nova Acta Reg. Soc. Sci. Upsal., ser. 4, 15(9). 92 pp., 70 figs. Tanaka, T. 1952. The systematic study of the Japanese Protoflorideae. Mem. Fac. Fisheries Kagoshima Univ., vol. 2(2). 92 pp., 41 figs., pis. 1-23. Thxjrkt, G. 1840. Note sur I'anthere du Chara et les animalcules qu'elle renferme. Ann. Sci. Nat. Dot., ser. 2, 14:65-72, pis. 5-8. 1850. Recherches sur les zoospores des algues et les antheridies des cryptogames. Premiere partie. — Zoospores des algues. Ann. Sci. Nat. Bot., ser. 3, 14:214- 260, pis. 16-31. 1851. Recherches sur les zoospores des algues et les antheridies des cryptogames. Seconde partie. — Antheridies des cryptogames. Ann. Sci. Nat. Bot., s^r. 3, 16:5-39, pis. 1-15. 1853a. Sur la fecondation des Fucacees. Compt. Rend. Acad. Sci. [Paris], 36:745-748. 1853b. Note sur la fecondation des Fucac6es. Mem. Soc. Sci. Nat. Cherbourg, 1:161-167. 1854. Recherches sur la fecondation des Fucacees, suivies d'observations sur les antheridies des algues. Ann. Sci. Nat. Bot., ser. 4, 2:197-214, pis. 12-15. 1855. Recherches sur la fecondation des Fucacees et les antheridies des algues. II. Ann. Sci. Nat. Bot., ser. 4, 3:5-28, pis. 2-4. 1875. Essai de classification des Nostochinees. Ann. Sci. Nat. Bot., ser. 6, 1:372-382. Thuret, G., and fi. Bornet 1878. Etudes phycologiques. iii + 105 pp., 51 pis. Paris: Masson. Thwaites, G. H. K. 1847. On conjugation in the Diatomaceae. Ann. Mag. Nat. Hist., 20:9-11, 343-344, pis. 4, 22. 1848. Further observations on the Diatomaceae; with descriptions of new genera and species. Ann. Mag. Nat. Hist., ser. 2, 1:161-172, pis. 11, 12. Tiffany, L. H. 1930. The Oedogoniaceae, ... 253 pp., incl. 64 pis. Publ. by author. Tilden, Josephine E. 1933. A classification of the algae based on evolutionary development, with special reference to pigmentation. Bot. Gaz., 95:59-77, pi. 1. 1935. The Algae and their Life Relations, xii + 550 pp., 257 figs., frontis. Minneapolis: Univ. Minnesota Press. TONI, J. B. DE 1888. Conspectus generum chlorophycearum hucusque cognitorum. Notarisia, 3:447- 453. 1889. Sylloge algarum . . . Vol. 1. Chlorophyceae. 12 4- cxxxix + 1315 pp. Padua: Publ. by author. 222 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES TONi, J. B. DE (Cont.) 1890. Osservazioni sulla tassonomia delle Bacillariee (Diatomee) seguite da un pros- pette dei generi delle medesime. Notarisia, 5:885-922. 1891a. Sylloge algarum . . . Vol. 2, Bacillarieae, Sec. I, Raphideae. cxxxii + 490 pp. Padua: Publ. by author. 1891b. Systematische Uebersicht der bisher bekannten Gattungen der echten Fucoi- deen. Flora, 74:171-182. 1892. Sylloge algarum . . . Vol. 2, Bacillarieae, Sec. II, Pseudoraphideae. Pp. 491-817. Padua: Publ. by author. 1894. Sylloge algarum . . . Vol. 2, Bacillarieae, Sec. Ill, Cryptorhaphideae. Pp. 819- 1556. Padua: Publ. by author. 1895. Sylloge algarum . . . Vol. 3, Fucoideae. xvi + 638 pp. Padua: Publ. by author. 1897. Sylloge algarum . . . Vol. 4. Florideae, Sec. 1. xx + Ixi +386 + [2] pp. Padua: Publ. by author. Transeau, E. N. 1951. The Zygnemataceae. xiv + 327 pp., incl. 41 pis. Columbus: Ohio State Univ. Press. Turner, D. 1802. A Synopsis of the British Fuel. Vols. 1 and 2. xlvi + 400 pp. London: White. 1808. Fuci . . . Vol. 1. 164 + [2] pp., pis. 1-71. London: Arch. 1809. Fuci . . . Vol. 2. 162 + [2] pp., pis. 72-134. London: Arch. 1811. Fuci . . . Vol. 3. 148 + [2] pp., pis. 135-196. London: Arch. 1819. Fuci . . . Vol. 4. [1] + 153 + [2] + 7 pp., pis. 197-258. London: Arch. TURPIN, P. J. F. 1828a. Apergu organographique sur le nombre deux. Mem. Mus. Hist. Nat. [Paris], 16:295-344, pi. 13. 1828b. Observations sur le nouveau genre Surirella. Mem. Mus. Hist. Nat. [Paris], 16:361-368, pi. 15. TUTTLE, A. H. 1926. The location of the reduction divisions in a charophyte. Univ. Calif. Publ. Bot., 13(12): 227-235, incl. pis. 22, 23. Vaillant, S. 1721. Caracteres de quatorze genres de plantes; le d^nombrement de leurs especes ; les descriptions de quelques-unes, & les figures de plusieurs. Hist. Acad. R. Sci. [Paris], Mem. Math.-Phys., 1719:9-47, 4 pis. Vaucher, J. P. £:. 1803. Histoire des conferves d'eau douce . . . xv + 285 + [2] pp., 17 pis. Geneva: Paschoud. 1821. Memoire sur les charagnes. Mem. Soc. Phys. Hist. Nat. Geneve, 1:168-179, 1 pi. ViLLAUS, D. 1789. Histoire des plantes de Dauphine. Vol. 3. xxxii + 1091 pp. Grenoble: Publ. by author. Vlk,.W. 1931. Uber die Struktur der Heterokontengeisseln. Beih. Bot. Centr., 48:214-220, 15 figs. 1938. Uber den Bau der Geissel. Archiv Protistenk., 90:448-488, 12 figs., pi 23. Wallich, G. C. 1858. On Triceratium and some new allied forms, with figures of the same. Quart. Journ. Micr. Sci., 6:242-253, pis. 12, 13. PAPENFUSS: CLASSIFICATION OF THE ALGAE 223 Wallicu, G. C. (Cont.) 1860a. Results of soundings in the north. Atlantic. Ann. Mag. Nat. Hist., ser. 3, 6:457-458. 1860b. On the development and structure of the diatom-valve. Trans. Micro. Soc, n. s., 8:129-145. 1861. Remarks on some novel phases of organic life, and on the boring powers of minute annelids, at great depths in the sea. Ann. Mag. Nat. Hist., ser. 3, 8:52-58, 7 figs. 1865. On the structure and affinities of the Polycystina. Trans. Micro. Soc. London, 13:57-84. 1869. Coccoliths and coccospheres. Quart. Journ. Micr. Sci., 17:77-78. 1877. Observations on the coccosphere. Ann. Mag. Nat. Hist., ser. 4, 19:342-350, pi. 17. Wallroth, F. W, 1815. Annus botanicus, sive supplementum tertium ad C. Sprengelii floram halensem. Cum tractatu et iconibus VI Charam genus illustrantibus. xxx + 200 pp., 6 pis. Halle: Kiimmel. (Not seen.) 1833. Flora cryptogamica germaniae. Pars posterior [=Vol. 2], continens algas et fungos. Ivi + 923 pp. Niirnberg: Schrag. Walz, J. 1866-1867. Beitrag zur Morphologie und Systematik der Gattung Vaucheria DC. Jahrb. Wiss. Bot., 5:127-160, pis. 12-14. Warming, E. 1876. Om nogle ved Danmarks kyster levende Bakterier. Videnskab. Medd. Dansk Naturhist. Foren. Kj0benhavn, 1875:307-420, pis. 7-10. 1890. Handbuch der systematischen Botanik. (Translated from the 2nd Danish ed., 1884, by E. Knoblauch.) xii + 468 pp., 573 figs. Berlin: Gebr. Borntraeger. Weismann, a. 1887. Ueber die Zahl der Richtungskorper und iiber ihre Bedeutung fiir die Verer- bung. viii + 75 pp. Jena: G. Fischer. West, G. S. 1904. A Treatise on the British Fresh-water Algae, xv + 372 pp., 166 figs., frontis. Cambridge Univ. Press. 1916. Algae. Vol. 1 [only vol. published], viii + [2] +475 pp., 271 figs. Cambridge Univ. Press. 1927. A Treatise on the British Fresh-water Algae . . . Rev. ed. by F. E. Fritsch. xvii + [1] + 534 pp., 207 figs., frontis. Cambridge Univ. Press. West, W., and G. S. West 1897. Welwitsch's African fresh-water algae. Journ. Bot., 35:1-7, 33-42, 77-89, 113-122, 172-183, 235-243, 264-272, 297-304, pis, 365-370. Wettstein, R. R. 1901. Handbuch der systematischen Botanik. Bd. 1. ix + [1] + 201 pp., 126 figs. Leipzig and Vienna: Deuticke. 1924. Handbuch der systematischen Botanik. 3rd ed. viii + 1017 + [1] pp., 653 figs. Leipzig and Vienna: Deuticke. WlLLE, N. 1890-1891. Conjugatae [und] Chlorophyceae. In A. Engler, Die natiirlichen Pflanzen- familien . . . Teil I, Abt. 2. Pp. 1-175, 128 figs. Leipzig: W. Engelmann. 1894. Ueber die Befruchtung bei Nemalion multifidum (Web. et Mohr) J. Ag. Ber. Deutsch. Bot. Ges., 12 [Suppl.] : (57)-(60), 6 figs. 224 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Welle, N. (Cont.) 1900. Algen aus dem nordlichen Tibet, von Dr. S. Hedin im Jahre 1896 gesammelt. Petermanns Mitt. Justus Perthes Geogr. Anstalt, Erganzungsband 28 (Heft 131), pp. 370-371. 1901. Algologische Notizen VII. Zur Verbreitung der Siisswasseralgen im siidlichen Noi'wegen. Nyt. Mag. Naturvidenskab., 39:1-22. 1909a. Algologische Notizen XV. uber Wittrockiella Nov. Gen. Nyt. Mag. Naturviden- skab., 47:209-225, pis. 11-14. 1909b. Conjugatae und Chlorophyceae. In A. Engler and K. Prantl, Die natiirlichen Pflanzenfamilien . . . Nachtrage zum I. Teil, Abt. 2. Pp. 1-136, figs. 1-70. Leipzig: W. Engelmann. Williams, J. L. 1904a. Studies in the Dictyotaceae. I. The cytology of the tetrasporangium and the germinating tetraspore. Ann. Bot, 18:141-160, pis. 9, 10. 1904b. Studies in the Dictyotaceae. II. The cytology of the gametophyte generation. Ann. Bot., 18:183-204, pis. 12-14. Williams, May M. 1925. Contributions to the cytology and phylogeny of the siphonaceous algae. I. The cytology of the gametangia of Coclium tomentosum (Stack.). Proc. Linn. Soc. New South Wales, 50(2) : 98-111, 42 figs. WiTTROCK, V. B. 1877. On the development and systematic arrangement of the Pithophoraceae, a new order of algae. Nova Acta Reg. Soc. Sci. Upsal., extra ord. vol. (Art. 19). 80 pp., 6 pis. Wood, R. D. 1952. The Characeae, 1951. Bot. Rev., 18:317-353. WORONIN, M. 1869. Beitrag zur Kenntniss der Vaucherien. Bot. Zeit., 27:137-144, 153-162, pis. 1, 2. Yamanouchi, S. 1906a. The life history of Polysiphonia violacea. Bot. Gaz., 41:425-433. 1906b. The life history of Polysiphonia violacea. Bot. Gaz., 42:401-449, 3 diagrams, pis. 19-28. 1909a. Mitosis in Fucus. Bot. Gaz., 47:173-197, pis. 8-11. 1909b. Cytology of Cutleria and Aglaozonia (a preliminary paper). Bot. Gaz., 48: 380-386. 1911. On the life history of Zanardinia collaris Crouan. (A preliminary note.) Bot. Mag. Tokyo, 25:9-11. (In Japanese.) 1912. The life history of Cutleria. Bot. Gaz., 54:441-502, 15 figs., pis. 26-35. 1913. The life history of Zanardinia. Bot. Gaz., 56:1-35, 24 figs., pis. 1-4. Zanardini, J. 1843. Saggio di classificazione naturale delle ficee. 65 + [1] pp., 3 charts, 1 pi. Venice: Publ. by author. Zaneveld, J. S. 1940. The Charophyta of Malaysia and adjacent countries. Blumea, 4(l);l-223, 21 figs., 16 tables. Zimmermann, W. 1930. Neue und wenig bekannte Kleinalgen von Neapel I-V. Zeitschr. Bot., 23:419- 442, 11 figs., pi. 1. ZiNNECKEB, EMMI 1935. Reduktionsteilung, Kernphasenwechsel und Geschlechtsbestimmung bei Bry- opsis plumosa (Huds.) Ag. osterr. Bot. Zeitschr., 84:53-72, 6 figs. MYCOLOGY By ERNST ATHEARN BESSEY Michigan State College, East Lansing The word "mycology," applied to the study of fungi, is not very many years older than the beginning of the hundred-year period covered by this series of papers. In the Latin form, mijcologia, it was used by Persoon (1801). As an English word, according to Murray (1908), it was first used by the Reverend Miles Joseph Berkeley in 1846 in British Flora, Fungi, in which, also, he applied the term "mycologist" to the students of fungi. In 1850 Fresenius used the word in the German form. After that it came into general usage in European publications in France and Italy, as well as in England and Germany, though in England the word "fungology" was frequently used, a term introduced by Berkeley in 1860. Fungi were known to the ancients. Indeed the Emperor Nero was very fond of eating the mushroom Amanita caesarea Schaff. ex. Fr., the specific epithet being given because of this fact. In the seventeenth and eighteenth centuries the larger fungi attracted the attention of botanists more and more but it was not until the publication of the works of Christiaan Ilendrik Persoon (1801, 1822-1828) and Elias Magnus Fries (1821-1832 and many subsequent publica- tions until about the time of his death in 1878) that the larger fungi were studied extensively as well as intensively. The microscopic fungi were mostly given scant attention or entirely passed over until the improvements of the compound microscope made it possible to study their structure and to begin to form systems of classification for them. The path-breaking work of Corda (1837- 1854) was scarcely completed by the middle of the last century. By the use of the microscope and the numerous illustrations in his great work, he added thous- ands of microscopic or semimicroscopic species to our knowledge. It must be remembered that one hundred years ago many botanists and other students of natural history believed that the small fungi occurring on or within the tissues of plants and animals were not distinct beings but actually modifica- tions of the diseased tissues of the host organisms, or "exanthemata." This was the view held by Elias Magnus Fries (1821-1832, 1836-1838), and Friedrich Wilhelm Wallroth (1833). In this same year Franz Josef Andreas Nicolaus linger, in one of the earlier books on pliytopathology, Die Exantheme der Pflan- zen (1833), supported these ideas. Twenty years later the English botanist, John Lindley, in his book The Vegetable Kingdom, altliough apparently ques- tioning the development of fungi by other means than from spores, asserts that many botanists still hold to the vieAvs of Fries and Unger. Yet he doubts the ability of fungi to cause plant disease, indicating that they enter tissues already diseased from other causes, such as extreme moisture, drought, malnutrition, and so forth. This whole question is very dramatically set forth by Large in his very [225] 226 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES PIER ANDREA SACCARDO 1845-1920 SIMON SCHWENDENER 1829-1919 ROLAND THAXTER 1858-1932 LOUIS RENE TULASNE 1815-1885 BESSEY: MYCOLOGY Til interesting book Advance of the Fungi (1940), especially with reference to the great epiphytotic of late blight of the potato. In 1845 when this attacked the crops of Great Britain and Ireland and other parts of Europe there were two opposed groups of scientists. One, headed by Berkeley, insisted that the fungus associated with the disease (later named PJiytopMhora infestans [Mont.] De Bary) was the real cause of the trouble, whereas the other group, led by Lindley, maintained that the disease came first and was due to soil or weather conditions or to the "running out" of the varieties. As for the ever-present fungus, some held with Lindley that it was simply growing upon the already diseased tissue, but was not the cause of the disease. Others agreed with Unger and Fries in considering the fungus the product of the diseased tissue. The Structure and Life History of Fungi In Great Britain Berkeley was for many years the leading student of fungi, including those whose study required the use of the compound microscope. He described many hundreds of species of hitherto unrecognized fungi and was the backbone of the group which maintained that many of the smaller fungi were actual parasites (in the present sense of the word) upon the hosts. On the Continent, after Corda's death in 1849, the study of the smaller fungi as well as of the structure of the larger fungi was carried on by Joseph Henri Le- veille (who lived from 1796 to 1870), and very many were carefully described and illustrated by him, even though he still maintained that they originated as exan- themata upon the host plants and were not really parasites. But the researches of Berkeley, Fresenius, and especially Montague (b. 1784, d. 1866) and Tulasne, rapidly brought the scientific world to abandon this idea. Persoon (1801) said, it is true, of some fungi, "Locus natalis . . . in plerisque parasiticus est ut pleraeque plantae aphyllae parasiticae sunt," but it is not certain whether he re- garded a parasite as we do as obtaining its nourishment at the expense of, and causing injury to, its host, or whether he used the term in the old classical sense of a person obtaining his meals at the table of another. Schleiden in the third edition of his Grundziige der Wissenschaftlichen Botanik (1850) took a midway position on the question. He did not regard the rusts and smuts as independent organisms but only as diseases of plants. On the contrary, the fungi which grew in the intercellular passages of their hosts and emerged through the stomata he considered true parasitic plants. His work was the leading botanical textbook in Germany and had great influence upon the ideas of students of mycology. However, since he did not publish descriptions of new species of fungi, it re- mained, apparently, without much influence upon mycological systematists, who did little in the way of careful intensive study of the structure and life histories of the individual fungi. This newer method of the study of fungi was undertaken in France by Louis Rene Tulasne (b. 1815, d. 1885) and his brother Charles (b. 1816, d. 1884). The former did the more intensive mycological study, the latter made the marvelously beautiful illustrations for their publications. It soon became apparent to them that some fungi had more than one type of spores and that these did not always germinate in a similar manner. In 1853 they demonstrated that spores of some rusts germinated by the formation of long hyphae or germ tubes and that others 228 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES produced upon germination hyphae of limited growth (which they called "pro- mycelia"), which bore usually four "sporidia." In 1854 came the discovery that the spores of the genus Uredo were only one stage of the development of a rust, the teleutospores {Puccinia, Uromyces, Coleosporium, Melampsora, etc.) being produced later by the same mycelium that had given rise to the uredospores. These rusts, therefore, had three spore forms, which we now call urediospores, teliospores, and sporidia (or basidiospores). The Tulasne brothers suggested that the absolute proof of this could only be obtained by controlled inoculation experiments which they had not made and which they believed would be very difficult to carry out successfully. In their Selecta Fungorum Carpologia (1861- 1865) — perhaps the most sumptuous work of the period, with illustrations whose artistic excellence has never been equaled — were demonstrated the various forms of reproduction of different fungi. It must be admitted that some of the vari- ous spore forms which the authors attributed to the fungi so beautifully and ac- curately illustrated were due to contamination by saprophytes or even parasites, which had nothing to do with the life histories of the fungi under study. Thus a pycnidial stage was described and illustrated for the Erysiphaceae, but later this was demonstrated by De Bary and Woronin (1870) to be a parasitic im- perfect fungus, Cicinnoholus, growing and producing its own pycnidia within the hyphae of the Erysiphaceae. Hence it became more and more evident that it was necessary to grow the fungi whose life history was under study from spore to maturity in pure cul- tures, free from the opportunity of access by other organisms. Due credit for the early making of cultures of fungi should be given to the Italian Pier' Antonio Micheli, who lived from 1679 to 1737. In his great work (1729) he described cultures on suitable vegetable media, using the spores of fungi that he called Mucor, Aspergillus, and Botrytis. The media were kept covered by bell-jars and developed only the fungus whose spores were sown upon them whereas similar pieces, not thus covered, developed "Mucor" Micheli's conclusion was that the spores of these various molds were normally distributed through the air. The pure-culture study method was in modern times first carried out suc- cessfully by Anton De Bary. He was born in 1831, the son of a busy physician in Frankfurt a. M., Germany. He obtained his M.D. degree at the University of Berlin at the age of twenty-two and immediately entered upon the practice of medicine in his home city. He admitted later that the diseases of his patients interested him only until he was sure of the correctness of his diagnosis, and so he soon gave up his practice, as he jokingly remarked "im Interesse der Kran- ken." In December, 1853, he became Privatdozent for botany in the medical faculty of the University of Tubingen. His biographer, Ludwig Jost (1930), states that he remained there only two years, "zweifellos iDeniger Kolleg lesend als forschend tdtig." He then accepted a call as Professor at the University of Freiburg, remaining there twelve years and gathering around him a coterie of eager students. In 1867 he was called to the University of Halle a. S. where he remained until his appointment in 1872 to the chair of botany at the newly founded University of Strasburg, a position that he held until his death in 1888. The botanical laboratory that he established at Freiburg in 1855 was one of the first half-dozen botanical laboratories in the world. He attracted students from many countries by his own boundless energy and by the inspiration which BESSEY: MYCOLOGY 229 impelled them to tireless research. By his scrupulous exactness of observation and teaching he gave to his students and to the readers of his published works a new appreciation of what fungi were and of their relationships. His first pub- lication, in 1852 on Achhja prolifera, was a result of research carried on during the last months before he took his final medical examinations and received his degree. This paper was followed in 1853 by a 144-page booklet entitled Unter- suchungen iiher die Brandpilze (including at that time the fungi now placed in the orders Ustilaginales, Uredinales, Protomycctales, and Peronosporalcs). Ac- cording to Jost, the chief points demonstrated in this second paper were the presence of a mycelium in all these groups from which, in definite ways, arose the characteristic spores. They were not the products of metamorphosed diseased host tissues. This was at a time when many botanists still had the idea that these fungi were the products of the transformation of the diseased tissues of the hosts. It was not until 1863 that De Bary wrote a paper in which he described the course of development of some Peronosporaceae from the formation of the conidia, their germination upon and infection of the host plants, the progress of the fungus in the host, and the formation of the asexual conidia and of the sexual organs, the oogones and antherids. He found the latter organs also in Eufotium and followed the development of the perithecium and asci and asco- spores. He also demonstrated that the mold laiown as "Aspergillus glaucus" was the asexual phase of Eurotium. The slime molds early attracted his attention (1858, 1859, 1862). He studied the growth of the Plasmodium, the formation of the sporangia and spores, the germination of the latter, the formation of the flagellate amoebae, and the origin of the Plasmodium. Because the life history of the vegetative phases of develop- ment was clearly more animal than vegetable, he changed the name of the group from Myxomycetes to Mycetozoa and boldly asserted that they belonged outside the vegetable kingdom and among the Protozoa. They completely lack mycelium and have a long amoeboid (or plasmodial) stage, hence cannot be placed in the fungi. Although later studies have fully confirmed the validity of De Bary's researches on this group, the majority of botanists have obstinately clung to the old idea that the slime molds are plants belonging to the fungi. Probably the zoologists are partly to blame for not welcoming with enthusiasm their trans- fer from the realm of botany to that of zoology. Most zoologists, it is true, accept them as animals, but all the important books on the slime molds treat them as plants. (Lister, 1925; Hagelstein, 1944; Martin, 1949.) De Bary now extended his inoculation studies to the rusts (1863, 1865). He inoculated bean plants. {Plmseolus vulgaris L.). He placed the teliospores of Uromyces in drops of water on their leaves, putting a bell-jar over the plant to prevent accidental contamination and to maintain the humidity of the air. The resulting infection showed first spermogonia and then aecia, but not the uredia or telia. When, however, he sowed the aeciospores in a similar manner on the same species of host, he obtained uredia and telia. Thus he proved, what some mycologists had suspected, that all five sorts of spores — sporidia, spermatia, aeciospores, urediospores and teliospores — were successive spore forms of the same rust. He could get no infection by using spermatia and made the suggestion that they were perhaps the male cells which, although still continuing to be formed, had lost their function. It must be remembered that it was not until 230 A CENTURY OF PROGRESS IN WE NATURAL SCIENCES more than sixty years later that J. H. Craigie (1927a, 1927b) demonstrated that the spermatia are really functional male cells. When De Bary sowed the telio- spores upon the wheat plants, however, he obtained no infection, although ure- diospores were effective. Eemembering the tradition among the peasants that barberry {Berheris vulgaris L.) caused the "blasting" of nearby wheat, he placed the teliospores from wheat upon the barberry leaves and obtained spermogonia and aecia. The mystery was solved. He coined the two terms to be applied to rusts: "heteroecious" for those that alternated on two kinds of not closely re- lated hosts and "autoecious" for those that could develop aecia and telia upon the same host. His study of other rusts demonstrated that there were some in which certain stages were lacking (e.g., aeciospores or urediospores or both), so that only spermogonia and telia occurred, whereas in Endophyllutn the ure- dia and telia were lacking and the aeciospores took over the function of the teliospores and germinated by means of a promycelium which bore sporidia. In his later studies De Bary sought for the sexual organs in the Ascomycetes, Mucorales, etc. ,To accomplish this he developed methods of growing the fungus from a single spore in pure culture on sterilized liquid or solid media. Oscar Brefeld, one of his students, learned these methods from him and improved upon his technique. He published a series of fifteen Hefte entitled Botanische Untersuchungen . . . (1872-1912) on various fungi, from the Mucorales, yeasts, various other Ascomycetes, smuts, various other Basidiomycetes, etc. These show great mastery of the methods but reveal that he missed the basic underlying principles taught by De Bary, viz., that these techniques were to be used to dis- cover the facts from which the unbiased conclusions could be drawn. Thus De Bary had clearly shown that sexual reproduction did occur in some Ascomy- cetes, as he had also demonstrated it in various species of Saprolegniales, Pero- nosporales, and Mucorales, although in many of these fungi he showed that there was a tendency toward the occurrence of apogamy or parthenogenesis, with the partial or complete loss of the sexual organs. He considered this a downward modification. Brefeld, on the contrary, developed the hypothesis that there was no sexuality in the Ascomycetes or Basidiomycetes. With this in mind he made his cultures to prove the correctness of the hypothesis. When Brefeld did ob- serve what De Bary considered to be sexual organs, he claimed that they had no sexual functions. It must be said, in excuse for his error, that he was so suc- cessful in growing his fungi from single spores that he missed the demonstration that would have been convincing, had he mated his cultures of opposite sexual phases. He claimed that there were two evolutionary tendencies that had led from the Phycomycetes to the higher fungi. In both of the lines, sexuality was supposed to have disappeared. The asci in the Ascomycetes were, in his opinion, modifications of the sporangia or zoosporangia while the basidia of the Basidio- mycetes were modifications of the conidiophores of those Phycomycetes that pro- duced conidia instead of sporangia. In both these lines he postulated a reduc- tion of the number of spores from indefinite to eight or four in the Ascomycetes or from indefinite to four in the Basidiomycetes. The genera of the former class in which the asci produce many spores, e.g., Ascoidea, Theleholus, and Monascus, he placed in the intermediary group, Hemiasci. It must be noted that later studies of Monascus demonstrated that this actually produces many eight-spored asci, tlie dissolution of whose ascus walls sets the ascospores free within the peri- BESSEY: MYCOLOGY 231 thecium, so that Brefeld erroneously thought that there was only a single ascus with many spores. Similarly, he postulated an intermediate group, Hemiba- sidii, for the Ustilaginaceae in which the promycelium is several-celled and pro- duces a variable number of sporidia. This was considered to be an early step toward the promycelium of Tilletia, in which there is only one cell and a smaller (but rather variable) number of sporidia is borne at its apex. From that to the Eubasidii, with normally four basidiospores at the top of the one-celled basi- dium, was the next step. When De Bary criticized these ideas of Brefeld, the latter became bitter and finally began to claim for himself the pure-culture method of the study of fungi (although in his first publications he credited his revered teacher with its invention). It is interesting that, although Brefeld's contention that sexual reproduc- tion was entirely lacking in the higher fungi was long ago disproved for Asco- mycetes and Basidiomycetes (Harper, 1896; Dangeard, 1907), his system of classification, modified to be sure, has long held sway in Germany and elsewhere and was retained in the revised edition of Engler and Prantl in 1928. The lichens were not studied as intensively by De Bary as the other fungi. Yet because of the similarity of their "gonidia" to free-living algae he suggested (1866) two possibilities as to their function in the lichen: either the mature lichens were the completely developed fruiting conditions of organisms ("goni- dia") whose incompletely developed forms were placed among the algae as Nos- tocaceae, etc., or they are typical algae which had become parasitized by certain fungi of the Ascomycetes. The latter suggestion may well have been what led Simon Schwendener to his interpretation of the role of the fungi and algae in the lichens, which he demonstrated in 1867 and 1868. Friedrich Wilhelm Zopf (b. 1846, d. 1909) was for many years Professor of Botany at the University of Halle a. S. He made extensive studies on the Chy- tridiales and other small aquatic fungi parasitic in algae and small animals. His textbook on fungi (1890) was, next to that of De Bary, the outstanding work on the subject for many decades. I must not fail to call attention to the very extensive mycological work done by the Englishman, Dr. A. H. Reginald Buller, who was for the greater part of his mycological career Professor of Botany at the University of Manitoba. His student work was carried on in England, where he received the B.Sc. and D.Sc. degrees, and at Leipzig, where he obtained the Ph.D. degree. Thus he combined in his training the best of the British and German traditions. His major studies were reported in a series of seven volumes entitled Researches on Fungi (1909- 1950). These contain detailed reports of his very ingenious experiments and careful observations on the activities and structures of fungi, mainly on Ure- dinales, Polyporales, and Agaricales, but including also Piloholus among the Mucorales, spore dispersal in the Ascomycetes and other fungi, etc. Besides these seven volumes he published numerous shorter notes of great interest, many of them in the British journal. Nature. Many of Buller's students have become prominent mycologists in Canada and the United States. In the foregoing pages I have omitted mention of the studies upon the fungi that attack man and other animals. Some of the forms that attack insects and form external fruiting bodies, e.g., Cordyceps, Isaria, etc., were described over two hundred years ago. At first there was a tendency to consider that the ap- 232 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES pearance above ground of the clavate stroma of Cordyceps, emerging from a caterpillar or other buried insect, was but a further development of the insect comparable to the metamorphosis of a pupa to a moth or butterfly. As early as the time of Persoon (1801) and Fries (1821-1832) the clavate stromata were recognized as fungi growing from within the dead insects. Mycotic growths in the air passages of birds were reported between 1815 and 1830. In 1837 Remak (according to Sartory, 1920) reported that in the diseases of man known as thrush and favus the whitish growth was a mass of fungus threads, an obser- vation confirmed two years later by Schoenlein and in 1841 by Gruby (Sartory, 1920). A number of other investigators reported similar discoveries in man and other animals in the next few years. In 1853 appeared the first collective work on these fungi, by Robin. Virchow (1856) described several cases of fungus- infection in the lungs of people and introduced the word mycosis for such infec- tions. From 1860 onward many different mycoses were reported, but mainly this was done by i)hysicians who had little mycological training. It was mainly among the French investigators in the next thirty to forty years that the great- est progress was made in medical mycology. R. Sabouraud (1894a, 1894b, 1910) made intensive mycological as well as clinical studies of the diseases caused by fungi attacking the hairs in man and other animals — the so-called tineas, ringworms, favus, and so on. E. Bodin (1901), Fernand Gueguen (1909), A Sartory (1920-1923), and Vuillemin (1931) wrote books bringing up to date the accumulated information on these diseases. In Germany, "Wilhelm Zopf (1890) devoted a considerable portion of his textbook on fungi to these parasites of man and other animals. In the United States, Dr. Carroll W. Dodge (1935) published a very extensive and detailed work on the subject, probably the most complete up to the date of its publication. Still more recent and clinically more modern is a book by Conant et al. (1945). It must be recognized that, except in the last two publications, the mycological nomenclature used is mainly that employed by medical writers, not actually in full accordance with the international rules of botanical nomen- clature. Vuillemin admits this in his discussion of the fungi attacking hairs, the "Trichophytes." In recent years the American students of medical my- cology have attempted to grow these fungi on standard culture media under, as far as possible, the same conditions of temperature, light, oxygen supply, etc., as are generally used foi? the culture of plant saprophytes. Thus it has become possible to determine the relationships of a number of these fungi, which, when grown on the special media and at 37° C, produced growths that did not at all reveal their kinship. It is not only in France and the United States that the study of medical mycology has been progressing. Very much has been accomplished in South America, Italy, Germany, Japan, and in other countries. The Taxonomy of Fungi While all the above-mentioned life-history and anatomical studies, as well as the special studies in medical mycology, were being carried on taxonomy of fungi was not neglected. The earlier European botanical writers included the larger nonmicroscopic fungi in their herbals, but with little idea of their real nature. BESSEY: MYCOLOGY 233 > . » MILES JOSEPH BERKELEY 1803-1889 OSCAR BREFELD 1839-1925 MORDECAI CUBITT COOKE 1825-1914 HEINRICH ANTON DE BARY 1831-1888 234 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES But with the opening of the eighteenth century there were a few students who gave great care and much time to the study and naming of fungi. Chief among these were Dillenius (1719) and Micheli (1729). These lived respectively from 1687 to 1747 and from 1679 to 1737. Linnaeus (1753) brought together in his twenty-fourth class of plants, Cryptogamia, the fungi whose names and descrip- tions he had found in the works of his predecessors but, since he knew little about fungi himself, the main value of this portion of his book was the application to these fungi of the binomials instead of the polynomial names of Micheli and Dillenius. After Linnaeus, the most significant mycological works in the next one hun- dred years, from the taxonornic viewpoint, were those of Persoon, Fries, and Corda. Christiaan Hendrik Persoon was born in South Africa in 1762. At the age of twelve he was sent by his father to Europe for his education. He never returned to Africa, although he kept in touch with his family and never lost his love for his fatherland. He studied in Holland and Germany and later went to France where he remained until his death. A very interesting account of the ancestry and life of Persoon is given by J. L. M. Franken (1937). The classification of fungi that he used in his Synopsis Methodica Fungorum (1801) and his Mycologia Europaea (1822-1828) was the foundation upon which the later mycologists based their work. The number of recognized genera and spe- cies had been greatly increased. The improvements of the microscope, although it was still a rather crude instrument, made it possible to study the manner in which the spores are borne; thus the fungi could be divided into major groups, many of which are still recognized. It must be remembered that by the Inter- national Rules of Botanical Nomenclature the Synopsis 3Iethodica is the authori- tative work for the names up to 1801 of the Uredinales, Ustilaginales, and Gas- teromycetes. Many mycologists believe that it would have been wiser to make that the date of reference for all fungi instead of using Linnaeus (1753), for the Mycetozoa and lichens, and Fries (1821-1832) for the rest of the fungi. Probably the work of Elias Magnus Fries (b. 1794, d. 1878), especially his Sy sterna Mycologicum (1821-1832), along with the above-mentioned publica- tions of Persoon, is what gave the great impetus to the taxonomic study of the larger fungi. For the next one hundred years the classification of the Agari- cales and Polyporales especially came to be based upon Fries. One must not for- get, however, that he in turn was dependent upon the clarity of vision of his predecessor, Persoon. Fries did not depend greatly upon the microscope so his knowledge of the smaller Ascomycetes and Fungi Imperfecti was not too good. August Carl Joseph Corda, who lived to be only forty (b. 1809, d. 1849), pub- lished a six-volume work, Icones Fungorum (1837-1854), in which he described and illustrated hundreds of microscopic fungi, using for that purpose a micro- scope that we would refuse to consider worth our while but which was good for his time. With the works of Persoon, Fries, and Corda the botanists inter- ested in fungi had at least a fair foundation upon which to build and a begin- ning of an idea of the structural features basic to taxonomy. It must be noted that among the foregoing authors there was considerable confusion as to what was meant by the terms "ascus" and "basidium." Appar- ently Fries did not distinguish between the "ascus" of the genus Agaricus and of Peziza. He criticized severely the emphasis of differences which could BESSEY: MYCOLOGY 235 not be distinguished except by the use of a microscope. It was not until about one hundred years ago that the ascus was clearly recognized as the cell within which the spores were produced, whereas the basidium had the spores external, or (as Fries considered it) extruded, from the apex of the "ascus." Indeed for many years the word "basidium" was used in a double sense : in the way we now use it as the structure upon which the basidiospores are borne (Berkeley, [1860] used it in this sense) ; or synonymously for a conidiophore, bearing a cluster of conidia at the apex, which is the usage in the earlier volumes of Saccardo's Sylloge Fungorum for the conidiophores or sporophores, as they were called later, of the Sphaeropsidales and Melanconiales. It was not until the publica- tion of Volume XXII of the Sylloge in 1913 that the change to these latter terms was made. A century ago the Reverend Miles Joseph Berkeley (b. 1803, d. 1889) was the leader in taxonomic mycology in England and indeed in almost the whole world. He wrote nearly four hundred papers on mycological topics and gave names to approximately six thousand new species of fungi. As noted previ- ously, at a time when most mycologists considered the microscopic fungi grow- ing upon or within plants to be merely "exanthemata" and not independent entities, he boldly maintained that Botrytis infestans Mont, (now known as Phytoplithora infestans) was the actual cause of the terrible potato disease which caused so much misery and death in Europe, especially in Ireland, in the mideighteenth century. He saw clearly the close connection that ought to exist between "vegetable pathology" and mycology. An account of his life and work, especially in reference to plant diseases is given by Knorr in Phytopatho- logical Classics, No. 8. Among his books may be mentioned Introduction to Cryptogamic Botany (1857) and Outlines of British Fungology (1860). Berkeley's successor in the study of fungi in England may be said to have been Mordecai Cubitt Cooke who lived from 1825 to 1914. He wrote the Hand- hook of British Fungi (1871), Handbook of Australian Fungi (1892) and nu- merous contributions to scientific journals. Perhaps his greatest service was the establishment of the periodical Grevillea, of which he was the editor and chief contributor for twenty volumes, from 1875 to 1892. Contemporaneous with part of Cooke's life and mycological activity were Worthington G. Smith and George Edward Massee (b. 1850, d. 1917). The latter was the first president of the British Mycological Society, one of the most valuable societies that has been established for the study of fungi. He was the author of British Fungus-Flora (1892-1895), A Textbook of Fungi (1910), European Fungus Flora, Agarica- ceae (1902), Monograph of the Myxogastres (1892), etc. Since then the number of fungus taxonomists in Great Britain has grown rapidly. It is impossible to mention more than a very few: Elizabeth M. Blackwell, Arthur Disbrowe Cot- ton, R. W. J. Dennis, Arthur and Gulielma Lister, E. W. Mason, Arthur A. Pearson, Thomas Petch, Carleton Rea, and Ethel M. Wakefield. The Commonwealth (formerly Imperial) Mycological Institute, in addition to functioning as a center for the plant pathology research of the Common- wealth, numbers among its staff workers who are carrying on a very large amount of taxonomic mycology of the highest excellence. Joseph Henri Leveille (b. 1796, d. 1870) was one of the outstanding mycolo- gists in France about one hundred years ago. He studied the nature of the 236 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES hymenium of the Hymenomycetes and introduced the term "basidium" in its present usage as long ago as 1837. He published in 1851 a report on the tax- onomy of the Erysiphaceae. Following his lead and that of the Tulasne brothers there arose in France many systematic mycologists. Space permits the naming of only a few: Philippe van Tieghem, Emile Boudier, Paul Vuillemin, Gabriel Arnaud, Herbert Bourdot, A. Galzin, Narcisse Patouillard, Julien Costantin, Henri Eomagnesi, Robert Kiihner, Roger Heim, Andre Maublanc, P. Konrad are among the many scholars who have brought honor to France. For many decades the Bulletin de lu Societe Mycologique de France, supplemented more recently by the Revue de Mycologie and many other periodicals, has published the contributions of these and other mycologists. It was to be expected that Germany and the other German-speaking lands of Central Europe would have many students of systematic mycology, although the impetus of De Bary's researches and teaching was strongly in the direction of anatomy and life history of fungi. In the early days of the hundred-year period under consideration we find the names of Rabenhorst, Fuckel, and Fresenius among these systematists. These were followed by Josef Schroeter, Gustav Lin- dau, Georg Winter, Eduard Fischer, Walter Migula, Andreas Allescher, Hein- rich Rehm, Paul Hennings, Paul and H. Sydow, Ernst Gauman, and a host of others. Four rather recent publications in the German language contributed greatly to the furtherance of the work in systematic mycology: Engler and Prantl, Die natUrlichen Pflanzenfamilien (1887-1938), Rabenhorst, Kryptoga- men-Flora von Deutschland, Oesterreich und der Schweiz (1884-1938), Schroeter, "Die Pilze Schlesiens" in Cohn, Kryptogamen-Flora von Schlesien and the great Kryptogamen-Flora der Mark Brandenburg by Lindau and others (1905-1938). Many of the mycologists listed above participated in the preparation of various portions of these works. Centers of Mycological Work In Italy the systematic study of fungi began very actively about one hundred years ago and has continued up to the present. In the first decade of the present century we find that Dorfler's Botaniker Adresshuch (1909) lists 307 Italian botanists, of whom 29 were noted as interested in mj^cology and 13 more in plant pathology. The outstanding student in this field was Pier' Andrea Sac- cardo (b. 1845, d. 1920). For a large portion of his active mycological career he was Professor of Botany at the University of Padua. He became interested in the fungi in the early seventies of the last century. He established the jour- nal Michelia in 1876 and continued its publication until 1882, when the great burden of writing the Sylloge caused him to cease piiblishing it. In Michelia appeared very many of Saccardo's first mycological contributions. Early in his mycological work he recognized that the descriptions of the fungi collected in all parts of the world were scattered far and wide, in all sorts of publications, such as local floras, monographs of genera, individual descriptions in various scientific periodicals, or even in agricultural or horticultural journals. For example, many of Berkeley's new species were described in Gardener's Chronicle. Furthermore, these descriptions were in various languages — Latin, Italian, Ger- man, French, English and others. Some were very brief, some very long drawn out. Thus it was impossible, unless a very extensive library was easily accessible BESSEY: MYCOLOGY 237 and the various languages were not serious barriers, to be sure of the identifi- cation of a fungus. After consulting with various other mycologists, and witli their encouragement and assistance, Saccardo entered (1882) upon the noted series of volumes entitled Sylloge Fungorum. In this magnum opus he attempted to bring together in systematic order all the published descriptions of fungi. Each description was in Latin and in a standard form, with the essential char- acters, including measurements, locality where found, etc. The work was planned to reach completion with Volume 8, which appeared in 1889, but the immense number of new species described in the meantime made it necessary to publish supplementary volumes. In the preparation of these later volumes, especially, he was assisted by various other mycologists, including his son-in-law, Alessan- dro Trotter; his son, Domenico Saccardo; Giovanni Battista Tra verso, Paul Sy- dow, and others. The last volume to appear (in 1931) was Volume 25, which brought up to date, as well as the disruption of World "War I and succeeding events permitted, all descriptions of fungi through 1920. It should be noted that the appearance of Volumes 22 to 25 was made possible in part by the active cooperation of Dr. W. G. Farlow, who interested various individuals and so- cieties in America in making available a considerable sum of money, to which Dr. P'arlow contributed. Since 1931, when Volume 25 finally appeared, the eco- nomic and political conditions have been such that it does not seem probable that further volumes will appear, at least not for many years. The consequence of the publication of the first and succeeding volumes of the Sylloge Fungorum was a tremendous upsurge in the description of new species whose authors had hesitated to describe them for fear of adding new names to species already described. Now it became possible for an investigator working far from an extensive library to venture to describe new fungi if his volumes of the Sylloge did not contain their descriptions. It must be admitted that not all mycologists were as modest as indicated above and that some of these kept rushing into print with "new species," regardless of the Sylloge. Because mycologists should at least know the names of new species and genera described since 1920 (i.e., after the publication of the last volume of the Sylloge) the Commonwealth Mycological Institute at Kew, England, has published, under the title Index of Fungi, lists of all such new species and gen- era or combinations from the year 1940 on. They have also collaborated in mak- ing available similar lists, prepared by Franz Petrak, for 1929 and 1932 to 1939. He is now working on material to fill in all the years from 1920 to 1940. Al- though these lists do not contain the descri])tions of these new fungi, they cite the place of publication so that mycologists may avoid duplication of names as well as know where to look for new species in genera in which they are interested. Saccardo is most widely known through the Sylloge Fungorum, but he was also the author of more than 140 lesser mycological contributions, including 14 numbers of Fungi veneti novi vel critici, from 1873 to 1882, and many numbers of miscellaneous contributions. His sporological systems of the Fungi Imperfecti (1880) and of the Pyrenomycetes (1876) exhibit the foundations upon which he based his classification of these groups. Aside from Saccardo and his collaborators in the preparation of the Sylloge there are many other Italians who .stand high in tlieir profession. Augusto Na- poleone Berlese is best known for his Icones Fungorum (1884^1905), but is the 238 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES JOB BICKNELL ELLIS 1829-1905 WILLIAM GILSON FARLOW 1844-1919 CHARLES HORTON PECK 1833-1917 NATHANIEL PRINGSHEIM 1833-1894 BESSEY: MYCOLOGY 239 author of many lesser contributions and of the more than 300-page monograph of the Peronosporaceae (1903), with many beautiful and accurate illustrations. Giovanni Battista Traverso was one of the leaders in the publication by the Societa Botanica Italica of the work Flora Italica Cryptogama (I, Fungi; III, Lichens) (1905-1938). Giovanni Bresadola lived most of his life in Trentino, while it was still in Austria, but he was an Italian by descent. His Iconographia Mycologica (1927-1932), one of the outstanding works for the fungi of south- ern Europe, was published in Milan by the Societa Botanica Italica. His Fungi Tridentini was published (1881, 1892) in Trieste. Other Italian mycologists are Teodoro Ferraris and Ralfaele Ciferri. Carlos Luis Spegazzini (b. 1858, d. 1926), of Italian nativity, settled in Buenos Aires and for several decades described hundreds of new species and a good many new genera of fungi. It may well be said that his work was the foundation upon which the knowledge of the rich mycological flora of Argen- tina and adjacent lands was founded. In such a vast country as Brazil with its varied climates, soils, and altitudes very much study still remains to be done on the systematic mycology of the republic. Professor Camillo Torrend pub- lished (1920-1935) a series of studies on the Polyporaceae of Brazil. Father Johann Rick, for many years a resident of the southernmost state of that country, Rio Grande do Sul, collected and studied the fungi. His interest was mainly in the Discomycetes, the larger Sphaeriales, the Thelephoraceae, and the Polypo- raceae (Rick, 1931-1936). The studies of the Brazilian fungi are now being pub- lished by Ahmes Pinta Viegas and A. Ribeiro Texeira, chiefly in the periodical Bragantia. For Venezuela and adjacent areas, aside from the studies by my- cologists from the United States and Germany, the most extensive publication is that by Chardon and Toro (1934). In Africa the main published work on systematic mycology in recent years in the Union of South Africa is that by Ethel M. Doidge of Pretoria and by P. A. van der Byl and Len Verwoerd of the University of Stellenbosch. From Uganda in the center of equatorial Africa we have extensive lists of fungi, including many new species, from the pen of C. G. Hansford, based upon extensive col- lections and studies made by him during his residence there. On the whole, however, the vast continent of Africa presents a mycological void. The Italians have published some lists of fungi collected by them from Eritrea and Italian Somaliland; and from Egypt Melchers (1931) published a check list of plant diseases and fungi, but that is from a rather limited area. In Asia the regions where active work in the study of the mycological flora has been carried on have been rather limited. In Japan, and more recently in China, in Ceylon, India, the East Indian islands, and the Philippines much has been done but very much more remains to be accomplished. The Russians have carried on quite extensive mycological explorations in Siberia and Russian cen- tral Asia, but that is so vast an area with such extremes of climate and vege- tation that only a good beginning has so far been accomplished. Tlie drier areas of southwestern Asia naturally have fewer fungi, but in the more humid val- leys separated by broad desert areas one would expect a high occurrence of endemism. A little work has been done by botanical explorers in Iran, and at present the botanists of Israeli and of Turkey are active, but they have as yet barely scratched the surface. The enormous high mass of Tibet, Afghanis- 240 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES tan, Chinese Turkestan, and the western and southwestern portions of China still await mycological study. Little has been done in Burma or the northern parts of Malaya and Indochina. The Arabian peninsula is almost untouched myco- logically. It is perhaps safe to say that there will not be many new mycological discoveries made there until conditions of life and travel are safer. The fungi of the British possessions of Ceylon and India were mostly studied by scientists sent out from Great Britain for considerable periods of time or by men like Berkeley, who remained in England and studied collections made by travelers in those regions. Dr. Edwin John Butler was the Imperial Mycologist in India from 1905-1919, returning to England to become the Director of the Imperial Mycological Institute. He published an important paper on the genus Pythium (1907), several papers on various rusts in India, and was a collabora- tor with H. and P. Sydow in a series of five numbers entitled Fungi Indiae Orientalis (1906-1916). With collaboration of G. R. Bisby he published Fungi of India (Butler and Bisby, 1931). In the last two or three decades there has been a great upsurge in mycological publications of good quality by students of Indian birth, among whom should be mentioned S. R. Bose, M. K. Patel, M. J. Thirumalachar, B. B. Mundkur and B. N. Uppal. In Ceylon Marshall Ward studied tlie disease of coffee caused by the rust Hemileia vastatrix B. & Br. After his return to England he was succeeded by Thomas Fetch, who remained in Ceylon a good many years. He studied the fungus flora very intensively. His report, published in collaboration with G. R. Bisby (Fetch and Bisby, 1950), lists 2,214 species of fungi from Ceylon. For a great many years the Botanical Garden at Buitenzorg, Java, has been a center of botanical research in almost every field of botany. Among the pub- lications issued there have been a good many mycological papers. The occupa- tion of Java by the Japanese, the subsequent fighting for the recovery of the island, and then the revolution which resulted in the establishment of a repub- lic have greatly interrupted the botanical work, although the Japanese did not harm the research laboratories. Dr. K. B. Boedijn has survived these disturb- ances and is still doing some mycological research. The chief periodicals in which the mycological papers from Java are found are Annates du Jardin Botanique de Buitenzorg, Bulletin du Jardin Botanique de Buitenzorg, and Reinwardtia. It has been in Japan that the chief mycological work in Asia has been car- ried on. For the Fhycomycetes may be mentioned the work of Yosio Tokunaga on the Chytrids (1933-1934); Hiroharo Indoh on the Blastocladiaceae (1940) and Leptomitaceae (1939); Masaji Nagai on Saprolegniaceae (1931, 1933); J. Hanzawa (1915), Yoshihiko Yamamoto (1930) and Momoji Yamazaki (1934) on the genus Rhizopus. Sanshi Imai published papers on the Helvellaceae (1932), on the Japanese Geoglossaceae (1934-1942), on the Clavariaceae (1929-1941) and on the Agaricaceae (1933). There have been extensive studies of the Ure- dinales, especially the series of papers by Naohide Hiratsuka (1927 to 1939). Seiya Ito (1909 to 1922) has also piiblished accounts of the fungi of this group. In recent years a few Chinese botanists, of whom F. L. Tai and Lee Ling may be mentioned, have been publishing the results of their studies upon fungi col- lected in China. The disturbed political and economic conditions in that great country in the last twenty-five years have been very discouraging to mycological BESSEY: MYCOLOGY 241 work. This may also be said of mycological work in the Philippines, where con- siderable work was done by American and European botanists, aided by very able students of Philippine birth; but the occupation of the islands by the Jap- anese in 1941 and the destruction of the centers of research put an end for many years to mycological studies. The situation has been much brighter in Australasia. The last four decades have seen the publication of some excellent contributions to the knowledge of the smuts, rusts, Polyporaceae, and Gasteromyeetes of New Zealand by G. H. Cunningham (1924 to 1950) as well as of the Polyporaceae and Gasteromyeetes of Australia, by the same author (1944, 1950). Daniel MeAlpine published a book on the fungi of Australia (1895) and one book each on the rusts (1906) and smuts (1910) of Australia. On the larger woody and fleshy fungi John Burton Cleland published a number of contributions, some alone (1934-1935) and some with the collaboration of Edwin Cheel (1914—1931) or of Leonard Rodway (1928-1929). Lillian Eraser (1933-1935, 1936) and Eileen E. Fisher (1939, 1950) have studied the sooty molds and related fungi of Australia. Thus it is apparent that systematic mycology has progressed far in Australasia in some of the important groups of fungi. Mycological "Work in North America In North America the earliest important contributions to the knowledge of the fungi of the country were made by the Reverend Lewis David von Schweinitz (b. 1780, d. 1834). He collected fungi extensively in North Caro- lina and in Pennsylvania and his two publications (1822, 1832) listed more than 2,000 species, many hundreds of which he described as new to science. He possessed a compound microscope, good for that period. He followed in the main the system of Fries. Accordingly the group called by him (1832) Aseo- mycetes included both Ascomycetes and Basidiomycetes as these terms are now used. His class Hymenomycetes included Discomycetes as well as Agaricales and Polyporales of the more recent classification. After the death of von Schweinitz in 1834 the chief botanical interest in this country for the next thirty years or more was in the collecting and naming of the vascular plants of the West, which was rapidly being explored and set- tled. However, there were three botanists who maintained the interest in fungi during this period. The Reverend Moses Ashley Curtis (b. 1808, d. 1872) lived the greater part of his life in North Carolina (see Shear and Stevens, 1919). He became interested in the lichens in the late 1830 's and was for years in close correspondence with Edward Tuckerman, to whom he sent many specimens with full notes. In the mid-1840's he began a correspondence with M. J. Berke- ley that lasted until his death. He sent several thousand specimens of fungi to Berkeley, always with careful data. Many of them were described as new spe- cies with the authority given as "B. and C." The two published a joint contri- bution (1850-1854) upon the fungi in the herbarium of von Schweinitz which had come into the possession of the Philadelphia Academy of Science. Curtis ex- changed specimens freely with Michener, Ravenel, and other botanists. The larger portion of his herbarium is now in the British Museum but a good many of his specimens are in the Farlow Herbarium of Harvard University. Ezra 242 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Michener (see Shear and Stevens, 1917) was a contemporary of Curtis but lived longer (b. 1794, d. 1887). He was particularly interested in the plants of south- eastern Pennsylvania and made extensive collections, especially of fungi, of which he listed some 1,200 species from Chester County. He sent many of his fungi to Berkeley, some of which the latter described as new species. Perhaps Miche- ner's greatest contribution to mycology was his rearrangement of the herbarium of von Schweinitz which was deposited in the Philadelphia Academy of Science, but in a sadly neglected condition. His own mycological collection is now in the Mycological Herbarium of the United States Department of Agriculture. The third of these almost contemporaneous amateur mycologists was Henry W. Ra- venel (b. 1814, d. 1887). He was born and lived most of his life in South Caro- lina. He collected enthusiastically, especially lichens, which he sent to Tucker- man, and fungi, of which he sent large numbers to Berkeley, who described many new species with the tag "B. & Rav." He published little on fungi but issued five centuries of Fungi Caroliniani Exsiccati between 1853 and 1860. Between 1878 and 1882, in collaboration with M. C. Cooke, he issued eight cen- turies of Fungi Americani Exsicatti. Another somewhat later botanist who developed great interest in fungi was Charles Horton Peck (b. 1833, d. 1917) who became the botanist of the New York State Museum at Albany, a position he occupied from 1867-1915. He wrote a series of Reports of the State Botanist from 1871 to 1913, including de- scriptions of numerous species of fungi, chiefly Agarics, with many colored il- lustrations. Many of the fungi described were new to science. Owing to the fact that he had to depend upon the descriptions, often very meager, of the European fungi and never had the opportunity to study their type specimens or the species growing wild in their type localities it is not to be wondered at that some of his identifications were erroneous. Sometimes he applied the name of a European species to a fungus that was really an American one, or the name he gave to a supposedly new species was in error because the species already had a name in Europe. In spite of these mistakes, unavoidable under the cir- cumstances, the result of his forty-odd j-ears of study of American fungi was the description and naming and preserving in the New York State Herbarium of numerous fungi. This collection has been available for study and reference to the later mycologists, who could thus verify their own work. Peck's work and collections have aided and inspired many mycologists, among whom may be mentioned George Francis Atkinson (b. 1854, d. 1918), Calvin Henry Kauff- man (b. 1869, d. 1931), Andrew Price Morgan (b. 1836, d. 1907), William Al- phonso Murrill (b. 1869), Alexander Hanchett Smith (b. 1904) and many more. Atkinson was a member of the Botanical Department of Cornell University from 1892 to 1918. His interests were broad. In systematic mycology, he worked in his later years mostly upon the Agaricaceae, especially the genus Amanita. As a teacher he led many graduate students into the field of mycology. Kauffmann was connected with the Botanical Department of the University of Michigan from 1904 until his death in 1931. He published many papers, chiefly on Agaricaceae, including monographs of the United States species of the genera Arjnillaria (1923), Inocyhe (1924), GompMdius (1925a), Lepiota (1925b), and Clitocyhe (1926). His 7nagnum opus was the Agaricaceae of Michi- gan (1918). Among the many mycologists who were at one time for longer or BESSEY: MYCOLOGY 243 shorter periods his students were A. II. Smith, Edwin Butterworth Mains, j\Ia- rion Lee Lohman, Delbert Swartz, Bessie Bernice Kanouse, Adelia McCrea, Dow Vawter Baxter, Lee Bonar, Frank Boyd Cotner, Lewis Edgar Wehmeyer. William A. Murrill worked at the New York Botanical garden from 1904 to 1924, as assistant curator and curator of the mycological herbarium. He was the instigator, and from its first number until 1924 the editor, of Mycologia, which was the successor to, but not connected with, the Journal of Mycology. The Journal ceased publication upon the death of Professor W. A. Kellerman, its editor. In addition to his curatorial and editorial duties Dr. Murrill wrote many articles for Mycologia and for other journals, mostly upon the Polypora- ceae and Agaricaceae. He also wrote several local floras of the Agaricaceae (1912, 1911-1918)^ He wrote most of Volume 9 and part of Volume 10 of the North American Flora (1907-1916) including most of the Polyporaceae and the Boletaceae and part of the Agaricaceae. In addition he wrote upon the resu- pinate Polyporaceae (1920-1921, 1942). Since his retirement from the New York Botanical Garden he has carried on mycological studies for a number of years on the Agaricaceae and Boletaceae of Florida, in affiliation with the Her- barium of the University of Florida. IMurrill aroused much criticism becaiise of his breaking away from the Friesian tradition of generic limits, especially in the Polyporaceae, following in part the suggestions of P. A. Karsten (1879, 1882), in dividing the bulky genera into numerous smaller, more compact ones based upon color and various anatomical and chemical characters that Fries did not consider important enough to warrant making generic distinctions. It is true that not all Murrill's ideas have been universally adopted, but some modern mycologists such as Singer (1949), Bondarzew (Bondarzew and Singer, 1941), William Bridge Cooke (1940), A. H. Smith (1938) go even further; in the writer's opinion, correctly. The more conservative systematic mycologists for the greater part of a cen- tury, out of their great respect for Fries, did not venture to divide the single large genus Agaricus into smaller genera until Fries, himself, began to make this division. Lucien Quelet, in France (1872-1875), first used most of the Friesian subgenera as genera and Karsten (1879, 1882), following in the same line, added a good many more. As a result of the work of these mycologists and of others, between 240 and 250 genei-a of Agarics are now well defined, though not yet fully acknowledged. From the Friesian genus Polyporus have been produced in much the same way 40 to 60 genera. M. A. Donk of the Netherlands has followed along these lines in his studies of the Hymenomycetes of that country, bringing the nomenclature up to date (1928, 1931, 1933). Two names that have become established in connection with systematic my- cology in the United States are those of Job Bicknell Ellis (b. 1829, d. 1905) and Benjamin :\Iatlack Everhart (b. 1818, d. 1904) ; we find the familiar E. & E. appended to descriptions of hundreds of new species. Ellis became interested in fungi by entering into correspondence with Kavenel, a correspondence which continued until the latter's death. Ellis' earlier collections of fungi, beginning about 1870, were sent partly to Berkeley and partly to Cooke and a large num- ber of species are accordingly tagged B. & E. and C. & E. As his knowledge of fungi increased, he began describing new species independently. In 1875 he began the distribution of centuries of exsiccati entitled North American Fungi of which 244 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES twenty-five centuries were prepared, these being followed by Fungi Columhiani. In 1880 he became associated in his mycological work with the well-to-do amateur botanist, Everhart, with whom he published many articles and described hun- dreds of new fungi. In 1892 they published jointly a very fine book which is still of great value. The North American Pyrenomycetes, with excellent illus- trations by F. W. Anderson. In 1886, in conjunction with William Ashbrook Kellerman (b. 1850, d. 1908), Ellis and Everhart founded the Journal of My- cology, in which numerous articles of systematic mycological interest appeared, mainly under the authorship of the founders, singly or collectively. In 1889 this journal was taken over by the Section of Vegetable Pathology of the United States Department of Agriculture, which published it quarterly for three vol- umes until 1894. Then, after eight years of suspension. Dr. Kellerman took over the task in 1902, beginning with Volume 8 and continuing until the close of Volume 13, when his death pat an end to the publication. Centers of Advance A student of the history of any science soon notices that the progress of the subject is not an even one geographically but that the centers of advance are scattered here or there. Closer examination reveals that these locations are de- termined by the residence at those places of some one man or group of men who are enthusiastically studying and teaching the subject. Thus in Sweden many able students gathered around Linnaeus two hundred years ago. Eighty years later Fries had many followers. Around De Bary from 1853 until his death in 1888 there was always a group of zealous students. Farlow at Harvard seventy-five years ago began to attract men in the same way, and following him were Thaxter, Weston, and White, not to mention the many students trained there and going elsewhere to form centers of their own. This sort of propagation from old center to new centers is of course only possible to any considerable extent when the scholars at the centers are associated with colleges or univer- sites. So men like von Schweinitz, Curtis, Eavenel, Michener, and Ellis, al- though performing great amounts of excellent mycological work, could not prop- agate the spirit so widely as the men at Harvard, Cornell, Michigan, and other institutions. Coker and Couch form a mycological center at the University of North Carolina from which a good many mycologists have gone out. At Pur- due University the influence of Joseph Charles Arthur built up a group, scat- tered among various other institutions, of specialists in uredinology. At the University of Minnesota, under the influence of Edward Morse Freeman and Elvin C. Stakman, there are gathered men studying the various races of cereal rusts in their relation to their hosts and experimenting with the breeding of strains of rusts, as well as of hosts resistant to them. Often a sharp distinction cannot be made between the mycological and phy- topathological aspects of the subject. Thus in the study of the genus Fusarium, as carried out by Sherbakoff, Wollenweber, Hansen, Snyder, and others, the pathogenic activities of the strains under study must be considered along with the cultural and morphological characters. Thus it is that mycological work is apt to be found where there is also active phytopathological work. In recent years a new and very important branch of mycology has developed. BESSEY: MYCOLOGY 245 the study of the fungi that produces antibiotics. In my work as a plant patholo- gist I had occasion frequently to work with Petri-dish cultures of bacteria or of certain fungi. Occasionally my cultures were contaminated by the entrance of spores of a Penicillium or Aspergillus. Very often around such a contaminant the bacteria or fungi under culture were suppressed. I lacked the scientific curiosity to try to learn why it happened. I was not alone in my stupidity; I have talked with others who had the same experience. But there was one man, an Englishman named Alexander Fleming, who noted the destruction of the cells of a Staphylococcus around the contaminating colony of a species of Peni- cillium. He did not throw away the contaminated culture or cut out the invader while the colony was still small. He wanted to find out what was happening, and why. That is how penicillin was discovered. If the rest of us had been as keen as Fleming, penicillin could have been discovered decades earlier, for Na- ture had given us the opportunity to observe this phenomenon. Even though Fleming recognized the possible value of penicillin and tested it against various pathogenic bacteria it was not until his Penicillium notatuyn Westl. was studied by Florey and Heatley at Oxford University and sufficient penicillin was pro- duced to permit clinical experiments on human beings that the danger was past that this observation might be dropped from sight as merely an interesting fact. But with the outbreak of World War II a cooperative project was estab- lished in the United States, in which Florey and Heatley took part. Thus, as shown by Kenneth B. Eaper in his presidential address before the Mycological Society of America in 1951 (Raper, 1952), this international cooperation in- volved discovery of improved methods for more production of penicillin and development of improved strains of the fungus. So in the ten-year period from the beginning of this project the monthly production of penicillin in America, measured in "penicillin units," rose from 400,000,000 in May, 1941, to "between 23 and 33 trillion units" (23,000,000,000,000 and 33,000,000,000,000) ten years later. The success of this cooperative project with the product of Penicillium notatum started hundreds of investigators, independently and working for phar- maceutical manufacturers, to test thousands of cultures of all sorts of fungi (in- cluding Actinomycetes) and bacteria. The result is that more than three hun- dred antibiotics have been discovered, of which about seven are now in mass production and use. The search still goes on. The interested reader is referred to the ponderous work of Florey et al. (1949), in addition to this sketchy outline. Periodicals One hundred years ago there was not a single scientific periodical devoted solely to the publication of mycological contributions. Levcille published most of his important papers in Annales des Sciences Naturelles, Botanique, the ma- jority of De Bary's contributions appeared in BotaniscJie Zeitung of which he was the editor in the later years of his life. Among other scientific journals in which mycological papers appeared were Flora oder allgemeine Botanische Zei- tung; Pringsheira's Jahrhiicher fur Wissenschaftliclie Botanik; Berichte der Deutschen Botanischen Gesellschaft; Zeitschrift filr Botanik; Bulletin de la So- ciete Botanique de France, Comptes Rendus; Annals of Botany; Nature; Broteria; Nuovo Giornale Botanico Italico; Botanical Gazette; American Naturalist; Bui- 246 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES letin of the Torrey Botanical Club; Phytopathology; Botanical Magazine, Tokyo; Canadian Journal of Research; Tijdschrift over Plant enziekt en; Zeitschrift filr Pflanzenkrankheiten; Sve7isk Botanisk Tidskrift; and the annual transactions and bulletins of scores of scientific societies, academies, and other institutions. Possibly the earliest periodical devoted solely to mycology was Michelia, founded by Saccardo in 1876 and terminated in 1882. The Journal of Mycology was founded in 1886 and continued with some interruptions for fourteen volumes, ending in 1908. Mycologia began in 1909 and still continues actively. Transac- tions of the British Mycological Society began about 1916, the Review of Applied Mycology in 1922, the Bulletin Trimestriel de la Societe Mycologique de France, in 1895. Revue Mycologique began in 1879 and came to an end fifteen or twenty years later. Revue de Mycologie began in 1936. In Germany Mycologisches Cen- tralhlatt ran from 1912 to 1915, being then a casualty of World War I. An7iales Mycologici, 1903 to 1941, was a casualty of World War II, as was Zeitschrift filr Pilzkunde, founded in 1921. Sydowia was founded in 1947 as a sort of con- tinuation of Annales Mycologici. In Sweden Friesia ran for several years until the last war. Classification Systems The classification of fungi has naturally undergone great changes in the past one hundred years, corresponding with the increased knowledge of their struc- tures and life histories, on the one hand, and with the eventual general acceptance by mycologists of the hypothesis of evolution. Before the idea of evolution had gained acceptance, the degrees of relationships of plants (and animals) were based upon the greater or smaller degrees of similarity between the organisms that were being studied. The idea of "types" was proposed. These must not be confused with the nomenclatorial types of species, genera, families, etc., whose recognition is necessary to permit the application of the valid names upon these groups. In the older use of the term a "type" was an idealized organism, a sort of composite being, including the main characters of a large group of sup- posedly related plants or animals. Thus Ranunculus was the type of a whole group of Ranunculaceae, Magnoliaceae, Annonaceae, Berberidaceae, etc. These were all considered as having been created with va rious modifications of the type {Ranunculus in this case) — the greater the modifications, the less the degree of relationship. The idea was comparable to the work of an architect who draws a basic plan for a house and then modifies it in many ways so that, although the houses are basically similar, each one differs in a few or many particulars from the others. So the Creator was supposed to have formed his generalized type for a group of plants (or animals) and then, on the day of creation, to have modified this in some degree. It must be confessed that we who believe in evolution have had to take on some of the ideas of the earlier systematists. They measured the strength of what they called "relationship" by the degree of similarity, without accepting the idea of genetic kinshi]). We, too, use the degree of similarity to indicate the probable (or ])ossible) path of the evolutionarj^ change and so to indicate the degree of "blood relationslii])." As more fungi are studied and their struc- tures and life histories determined, we have become able to suggest what may BESSEY: MYCOLOGY 247 have been the more primitive forms and by what routes evolution may have pro- duced the different groups. Theoretically, now, the ideal system of classification will attempt to indicate these lines of descent (or shall we say, ascent) from the first organisms that we may call fungi. Since, however, fungi are not easily preserved as fossils, we cannot call upon the phytopaleontologists to assist us by showing what types of fungi occurred at each geological era. Therefore we have to depend upon the study of the ends of the twigs of the phylogenetic tree and by comparing these to surmise what the trunk and the main evolu- tionary branches probably were. Because of the structural differences in different groups of fungi and the different chemical constitution of their cell walls, as well as differences in their life histories, some of the earlier mycologists who believed in evolution concluded that the fungi are not necessarily a single phylctic series but that evolution from algae to fungi may have occurred at several different points. The necessary consequence of the acceptance of such a hypothesis would be belief in the poly- phyletic nature of the fungi we are acquainted with, in other words, these would not represent a great group of common descent. The different groups, arising from different algae, would not be interrelated, except as one traces relation- ship down through their various ancestral algal stocks to their common algal ancestor. Some of the suggested alga-to-fungus relationships are as follows : origin of Chytridiales (in the wide, older use of this term) from unicellular algae, taking into consideration the existence of certain somewhat intermediate genera which are still considered as algae but which live endophytically, e.g., Chlorochytrium, Endosphaera, RJwdochytrium, etc. On the contrary, it has been suggested that the Chytridiales are descended l)y simplification from Saprolegniales. Another suggested relationship is Vaucheria-Uke algae to Saprolegniaceae, taking into consideration the occurrence of the endophytic genus Phyllosiphon, showing that such an intermediate step may occur in this area of relationship. From the Saprolegniales would have arisen the Peronosporales and possibly, by sim- plification, the Chytridiales. The suggested origin of Monohlepkaris from Oedo- goniuyn is certainly erroneous, now that the structures and life histories of both have been more fully elucidated. Similarly, the supposed connection of Mucor and Spirogyra cannot be upheld. One hypothetical connection, Florideae to As- comycetes, suggested by Sachs (1874), has so many data to support it that to this day many mycologists, including the writer, are inclined to accept the hypothesis (see Bessey, 1942). The classifications of the pre-evolution days have undergone great modifica- tions, Fries (1821-1832) divided the fungi into four classes. 1. Coniomycetes: sporidia naked, without receptacles. Four orders, all except part of order Entophytae corresponding to our present Fungi Imperfecti. This latter order contained also the rusts and smuts. They were not true organisms, according to Fries, but exanthemata of diseased plants. 2. Hyphomycetes: thallus floccose, the sporidia borne upon or among the hyphae. These, too, were mainly Fungi Imperfecti. 3. Gasteromycetes: the whole fungus closed, containing the sporidia in its interior. This includes the present-day Gasteromycetes, the Mucorales, the Mycetozoa, and the Pyrenomycetes. 4. Hymenomycetes: hymenium soon exposed, bearing the sporidia superficially, in the 248 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES more perfect (i.e., typical) forms the sporidia included in the asci.i The class included sclerotial forms (Sclerotium, Ef^ysiplie, etc.); Tremella; Dacrymyces ; the Discomycetes; Solenia; Cypliella; with the highest order the Hymenini practically the same as the pres- ent day Hymenomycetes. Von Schweinitz (1832) follows Fries but rearranges the classes somewhat. A. Ascomycetes: bearing the sporidia in asci' Class I. Hymenomycetes asci on an open receptacle Class II. Pyrenomycetes, asci within perithecia B. Sporomycetes: bearing free sporidia, not in asci Class III. Gasteromycetes, sporidia free within a peridium Coniomycetes of Fries, sporidia without peridium Class IV. Hyphomycetes, sporidia borne directly on the thallus Class V. Gymnomycetes, sporidia borne on a sporodochium Berkeley (1857) made a considerable change in his classification of fnngi. By this time the studies of Montagne and Leveille had shown the difference be- tween the ascus and the basidium. The following is Berkeley's key. Fungales Sporidiiferi (sporidia in sacs) Ascomycetes: asci formed from the fertile cells of an hymenium Physomycetes: fertile cells seated on threads not compacted into an hymenium Sporiferi (naked spores) Hyphomj'cetes: spores naked, variously seated on conspicuous threads which are rarely compacted; mostly small in proportion to the threads Coniomycetes: spores naked, mostly terminal, seated on inconspicuous threads, free or enclosed in a perithecium Gasteromycetes: spores naked. Hymenium enclosed in a peridium, seldom ruptured before maturity Hymenomycetes: spores naked. Hymenium free, mostly naked, or if enclosed at first, soon exposed In the foregoing the Ascomycetes are the same as the group we now call by that name; the Physomycetes are practically identical with our Mucorales; the Hyphomycetes consist mainly of Fungi Imperfecti, but include also Perono- spora. The Gasteromycetes include, in addition to our present-day Gastero- mycetes, also the Mycetozoa; the Hymenomj^cetes include the Tremellales (in the wider sense), the Polyporales, and the Agaricales. The Coniomycetes in- clude Uredinales and Ustilaginales, in addition to some of the dematioid im- perfect fungi. The Saprolegniales are still included by Berkeley among the Conferva group of the algae, but with the doubt expressed that Achlya and its allied genera may be molds. In this connection it must be remembered that Nathaniel Pringsheim (1851, 1855, 1858, 1873) at first considered these fungi algae because their vegetative structures and manner of reproduction, sexual and asexual (the latter by zoospores), were in his opinion of greater weight in assigning them to a place in the algae than their lack of starch and chlorophyll. This seems to have been De Bary's opinion in his first paper on this group (1852). The next important classification of the fungi was that by De Bary (1866) in his textbook. He divides the fungi into four orders, the lowest, the Phyco- mycetes, coming first as revealing their more primitive nature and relationship 1. Remember that Fries, Schweintz, and other early mycologists did not set apart the basidia from the asci. BESSEY: MYCOLOGY 249 to the siphonaceous algae. From this order radiated the Hypodermii (smuts and rusts), the Basidiomyeetcs and the Ascomycetes, which he places highest in the fungal series. He has no group set aside for what we call the Fungi Imperfecta These he rather looks upon as asexual forms of Ascomycetes whose connections with the sexual stages have not been demonstrated. Eighteen years later De Bary (1884) modified this classification by establishing two series as follows. I. Ascomycetenreihe 1. Peronosporeen (nebst Ancylisteen unci Monoblepharis) 2. Saprolegnieen 3. Mucorineen Oder Zygomyceten 4. Entomophthoreen 5. Ascomyceten 6. Uredineen II. Von der Ascomycetenreihe divergierende oder der Steilung nacli zweifeltiafte Gruppen 7. Cliytridieen 8. Protomyces und Ustilagineen 9. Zweifeltiafte Ascomyceten (Saccliaromyces, etc.) 10. Basidiomyceten Groups 1-4, because of their near connection with the algae, are brought to- gether under the name Phycomycetes. In category II, groups 7 and 8 are to be treated in connection with the Phycomycetes, 9 naturally with the Ascomy- cetes, and 10 with 6 (Uredineae). The Lehrhuch der Botanik by Julius Sachs was of great influence in the de- velopment of botanical studies. This appeared in many editions and was trans- lated into several languages. In his earlier editions he followed De Bary for the classification of the fungi. In his fourth edition (1874) he adopted a quite different arrangement. He places the plants below the group Bryophyta in the group Thallophyta. This he divides into four classes, each containing plants with chlorophyll and those without it. The main line of evolution he indicates goes upward in the chlorophyll-containing series (i.e., the algae), the chloro- phyll-free organisms in each class being derived from those with chlorophyll in the same class. In other w^ords, the fungi are polyphyletic and do not form a single phylum. The four classes of Sachs are the following. I. Protophyta. No sexual reproduction Chlorophyll-containing Chlorophyll-free Cyanopliyceae Scliizomycetes (=Bacteria) Palmellaceae (in part) Saccliaromyces II. Zygosporeae. Sexual union of equal cells to produce a zygospore With chlorophyll Lacking chlorophyll Union of motile cells Volvocineae Myxomycetes (Hydrodictyeae) Conjugation of resting cells Conjugatae (including Diatomeae) Zygomycetes III. Oosporeae. Fertilization of egg to produce an oospore With chlorophyll Lacking Chlorophyll Sphaeroplea Vaucheria ( Saprolegnieae ) Peronosporeae Oedogonieae Fucaceae 250 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES IV. Carposporeae. Sexual reproduction results in the production of a spore fruit With chlorojihyll Lacking chlorophyll Coleochaeteae Ascomycetes (including the Lichens) Florideae Aecidiomycetes Characeae Basidiomycetes Sachs believed that the Saprolegniales and thence the Peronosporales arose from algae closely related to Vaucheria with a few differences : disappearance of chlorophyll, lack of free-swimming male gametes (these being replaced by a con- jugation tube from the antherid piercing to the egg), and the production of numerous simple biflagellate zoospores instead of a large compound zoospore with hundreds of pairs of flagella. The branched coenocytic vegetative struc- ture with cellulose cell walls, the production of zoospores in terminal segments of the hyphae, and the formation of large oogones with antherids usually aris- ing nearby are characters common to Vaucheria and the Saprolegniales. The idea that Mucorales represent developments from the Conjugatae in which the chlorophyll has been lost was adopted by Sachs from Brefeld, who emphasized the similarity of the formation of the zygospores in both groups of organisms. Although the suggestion of De Bary and Sachs that the Sapro- legniales are probably derived from Vaucheria-like algae has persisted in some quarters (Gaumann, 1949), mycologists have been led to reject the idea of the close relationship of these groups because of other factors: the type of hyphae tubular coenocytic in Mucorales, cellular with uninucleate cells, in Conjugatae; cell wall mainly of chitin in the former, of cellulose in the latter; and abundant production of asexual wind-borne spores in the former, no special asexual cells in the latter. In the fourth class, Carposporeae, the central feature is the production of a spore fruit, i.e., a mass of cells some of which are the spores which will produce the new plant. The spore fruit in the Ascomycetes, in which sexual organs are still functional, gives rise to ascospores contained in asci, at the ends of ascog- enous hyphae originating from fertilized oogones. Around these hyphae may also be present the vegetative hyphae that form the paraphyses and the main body of the perithecium or apothecium. In many of these Ascomycetes are pro- duced nonmotile spermatia which unite with the receptive threads (tricho- gynes) from the oogones and thus bring about the fertilization. This is similar to what happens in the Florideae and gave rise to Sachs's suggestion of the origin of the Ascomycetes from that group. This view still persists among many my- cologists (see Vuillemin, 1912, in which a very full discussion is given of the various suggested systems of classification of the fungi). Brefeld did not accept the ideas of Sachs and rejected those of De Bary, except the origin of the Oomycetes from the Siphoneae. For him, this group does not represent true fungi. The true fungi begin with the Mucorales, which he considers to have developed from algae that produced zygospores (e.g., Con- jugatae). He emphasizes that the algae retained their sexuality and evolved into the higher green plants. The primitive fungi (the Mucorales) quickly began to magnify the importance and complexity of their asexual reproduction at the expense of the sexual reproduction, which soon disappears as evolution pro- gresses toward the higher fungi. In the main line of fungus evolution, the basic group is class Zygomycetes (the class Oomycetes, in his view, comes to a blind BESSEY: MYCOLOGY 251 end). He divides this class into three series, leading to three lines of develop- ment. These are based on the asexual reproduction as follows : sporangia alone, sporangia and conidia, and conidia alone. The first group has fungi witli only the fully developed sporangia, as in Mucor, Rhizojnis, etc., or with both spo- rangia and sporangioles, as in Thamnidium. By reduction of the s]iorangioles to indehiscent, monosporous cells arose the conidia of the Choanephoraceae, forming the second group, in which true sporangia as well as these conidium- like sporangioles occur. Again from Thamnidium, by a similar reduction of the sporangioles to conidia and the complete loss of the true sporangia, came the third group of which Chaetocladium is characteristic. Tlie higher fungi, which Brefeld calls Myeomycetes, have entirely lost their sexuality. In the Ascomycetes the ascus is derived from the sporangium of some mold, like Choanephora and the conidia from the reduced indehiscent sporan- gioles. An intermediate group, the Hemiasci, is postulated, including fungi in which the sporangium, now well on its way to become an ascus, still remains with a large indefinite number of spores, the final step being the reduction of this large number of spores to a definite number, usually eight. From the com- pletely conidial Mucorales, such as forms of the same degree of development as Chaetocladium, Brefeld postulates the origin of the Hemibasidii, in which the conidiophore of this fungus has been reduced to a several-celled protobasidium with an indefinite number of spores. Here are the Ustilaginaceae and Tille- tiaceae. The former gives rise, with the number of spores reduced to four but with the basidium still several-celled, to the Protobasidiomycetes, including the Uredinales and AuricuJaria and Tremella and Pilacre. From the vicinity of Tilletia, with its one-celled promycelium or protobasidium, arose the Autobasi- diomycetes, with their spore number reduced to four. F. von Tavel (1892) de- votes a very interesting little book to a discussion of the fungi in the light of Brefeld's classification. Dangeard goes a step further in separating the fungi completely from the algae, thus forming two independent series. Both are assumed to have evolved from Protozoa of the group Flagellata. The algae became plants at the point of evolution where their flagellate, chlorophyll-containing ancestors lost the power of engulfing particles of food. The fungi arose from the flagellates that lacked chlorophyll, likewise at the point where they no longer took into their cells the particles of food. Thus the fungi are a kingdom parallel to the plant kingdom, on the one hand, and to the animal kingdom, on the other. It is worthy of note that G. W. Martin (1932) makes a somewhat similar suggestion. Wilhelm Zopf (1890) follows a classification similar in part to that of Bre- feld, but places the Ascomycetes last. In these he goes from the simple forms, like Saccharomyces, Endomyces, Gymnoascaceae, to, at the peak, the Pezizales. He recognizes the formation of ascogones in many Ascomycetes and even the union of these in Pyronema, with club-shaped "pollinodia," but expresses doubt as to their real sexual function. The classification of fungi followed in the first edition of Engler and Prantl (1897-1907) is, in its main features, the same as that of Brefeld. In the second edition (1926-1938) the main features are retained with some modifications made necessary by the cytological confirmation of the actual occurrence of sexu- ality in the higher fungi as well as in the Phycomycetes. Yet this fact has not 252 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES had the effect that it would seem to deserve, perhaps owing to the well-known conservatism of systematists and their reluctance to make changes in the familiar classification. So we find the monophyletic concept of the fungi strongly entrenched. To be sure, the ideas of Sachs and a number of others (as pointed out by Vuillemin, 1912), that the fungi have arisen from algae at various points, still persist. The most recent books to use the results of the recent cytological and anatomical studies in the classification are the two books by Ernst Albert Gaumann (1926, 1949). In his later book Gaumann recognizes four classes of fungi. The first class consists of the Archimycetes. They are endophytic parasites which, in their early vegetative stages, are naked and, in some cases at least, amoeboid. Later, the whole structure, often up to that stage still uninucleate, in spite of its growth, forms a cell wall and then by multiplication of the nuclei and division of the protoplasm becomes a zoosporangium or a gametangium, within which are contained the motile naked zoospores or gametes. Of the four families recog- nized two, Olpidiaceae and Sj'nchytriaceae, are usually placed in the Chytridiales, because of the posteriorly attached single flagellum. The other two families, Plas- modiophoraceae and Olpidiopsidaceae, have two anterior or lateral flagella and are usually placed respectively near the Myeetozoa and the Saprolegniales. These four divergent families are, according to Gaumann, probably to be assigned to an origin among the Flagellata. Gaumann's second class, the Phycomycetes is acknowledged to be at least diphyletic. The first three orders (Reihen), Chytridiales, Blastocladiales, and Monoblepharidales are certainly related. Their zoospores (and motile gametes, where formed) have a single posterior flagellum (which is of the whiplash type, as in the first two families of the Archimycetes) , and the cell wall does not typi- cally contain cellulose. They are derived from the Flagellata. The fourth Reihe, the Oomycetes, has cellulose-containing walls and the motile cells, where formed, have two anterior or lateral flagella, one of the tinsel type and one of the whip- lash type. The vegetative structure is a more or less branched, coenocytie hypha on which are formed rounded oogones, with one or more eggs (oospheres) which are fertilized by conjugation tubes from antherids that arise nearby or at a dis- tance and become attached to the oogone. The fertilized egg becomes a thick- walled oospore. This Reihe is so similar in structure to the Siphoneae, especially Vaucheria, that Gaumann seeks its ancestry in that general group, as did Sachs, De Bary, and others. It has several families, soil or water inhabitants and strict parasites in land plants. It ends blind, as the remainder of the fungi are not considered to have derived from the Oomycetes. The fifth Reihe is that of the Zygomycetes, from which the higher fungi are considered to have arisen. Vege- tatively they resemble the Oomycetes, in that they are branched tubular coeno- cytes, but their cell wall has chitin as its chief constituent. Sexual reproduction is by the union of two nearly equal and similar gametangia to form a thick- walled zygospore. Asexual reproduction is by the formation of sporangia, within which are produced the encysted spores, instead of the naked zoospores of the preceding class. These sporangia show great modifications, leading in several directions to the production of wind-borne conidia (which in most cases repre- sent indehiscent sporangia reduced in size, with contained spores reduced to one), BESSEY: MYCOLOGY 253 The third class, Ascomj^cetes, is considered to have arisen from some Zygo- mycete in which tlie union of two gametangia — instead of producing a zygo- spore, out of which later, on germination, a stalked sporangium arises — produces immediately the sporangium, which is specialized to become the ascus. In the subclass Protascales ascogenous hyphae and spore-fruits are lacking. Here come the orders Endomycetales (^ Saccharomycetales of authors) and Taphrinales. The subclass Euascomycetes includes the rest of the Ascomycetes, in which spore- fruits are built and the asci are produced on ascogenous hyphae. The basal group of the Euascomycetes is the order Plectascales, in which the ascogenous hyphae branch through the interior of the spore-fruit so that the asci are scat- tered throughout it. The asci and other tissues dissolve at maturity of the asco- spores so that the latter lie free in the now hollow ascocarp. The ascospores are not expelled from the asci. The spore-fruit shows varying degrees of complexity, from a loose weft of hyphae among which the ascogenous hyphae creep and pro- duce their asci (family Gymnoascaceae) to rather massive structures (e.g., Ela- phomycetaceae). Sexual reproduction by union of antherids with ascogones occurs frequently. From this order branch off two series of orders, the Ascolo- culares, in which the asci are formed in cavities which they dissolve out in the stromatic tissue, and the Ascohymeniales, in which these cavities are formed during the growth of the spore fruit and then become lined by a palisade of asci. The former group includes among other orders the Perisporiales, Myriangiales, Pseudosphaeriales. In the Ascohymeniales are found the Sphaeriales (including Hypocreales), Pezizales (=the operculate Discomycetes), Helotiales (inoper- culate Discomycetes) and Tuberales. The Laboulbeniales are placed at the close of the class with uncertain position as regards relationship. The fourth class, Basidiomycetes, is placed highest because of its derivation from the higher Ascomycetes (probably some of the Discomycetes). The basi- dium is looked upon by Gaumann as an ascus from which have emerged four exogenous pockets containing each a single ascospore. This ascospore with its containing wall is the so-called basidiospore. Gaumann recognizes two sub- classes: Holobasidiomycetes, with one-celled basidium, and Phragmobasidiomy- cetes, with the basidium longitudinally or transversely septate. The Holobasi- diomycetes he considers the more primitive type, derived from the Ascomycetes in which the hook or crozier (or its derived form, clamp-connections) is present. The Holobasidiomycetes are divided into the Hymenomycetes, in which the basi- diospores are violently expelled, and the Gastromycetes, in which the basidio- spores are passively distributed. In the second subclass, Phragmobasidiomycetes, the basidiospores are violently shot away in most of the species except in Family Ustilaginaceae. The following four groups are placed here : Tremellales, Auriculariales, Uredinales, and Ustilaginales. The late Herbert Spencer Jackson, for many years a student of the Uredi- nales, published a memoir (1931) in which he compared the life cycles of the rusts with those of the red seaweeds, suggesting that the similarities might indi- cate relationship between these groups. The spermogonium of the Uredinales may be considered to sliow relation- ship of the rusts to those Ascomycetes in which such structures occur. The ideas of relationship which the author has inherited and developed in the last half -century (see Bessey, 1942, 1950) may be outlined here. 254 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES The ]\Iycetozoa (Myxogastrales, Acrasiales, Labyrinthulales, and Plasmodio- phorales) are, following De Bary, placed outside the vegetable kingdom, being considered as derived from Protozoa of the group Rhizopoda and not progress- ing further to produce recognized fungus groups. When the flagella of the motile cells have been examined, they are found to occur in pairs, both of the whiplash type, thus barring any connection of tlie Plasmodiophorales with the Olpidiopsidaceae, where one of the flagella is of the tinsel- type and the other of the whiplash type. The true fungi are believed to begin with the Phycomycetes. The simplest of these fall into three series : Chytridiales, with a single posterior, whiplash type flagellum; Hyphochytriales, with a single anterior flagellum of the tinsel type; and Lagenidiales (including Woroninaceae and Olpidiopsidaceae, but not the Plasmodiophoraceae) with two anterior or lateral flagella, one of the tinsel type, the other of the whiplash type. The Chytridiales connect di- rectly with the Blastocladiales and Monoblepharidales, this line then ending blind; the Hyphochytriales have no recognized further development; the Lage- nidiales lead onward to the Saprolegniales and Peronosporales. Two paths of evolution of the Chytridiales, Hyphochytriales, and of the simpler Lagenidiales are suggested. They may be primitively simple, derived from some algal ances- tors of the group of Heterokostae, in which the flagella are of the two types. By the loss of the tinsel type flagellum the Chytrid type may have originated; by loss of the whiplash flagellum the Hyphochytrial group might have developed; while the Lagenidiales line may have had its beginning with the retention of both types of flagellum. But, contrariwise, these simple forms may have arisen by simplification from some fungi of the Lagenidiales, Saprolegniales, Perono- sporales lines which, as suggested by De Bary, Sachs, Gaumann, and others may have arisen from algae in the vicinity of Vaucheria. The author seeks the origin of the Mucorales in the soil-inhabiting Sapro- legniales in which the sporangia produce encysted spores (as in Ai^lanes) in- stead of the zoospores usually found in that order. The approximately equal gametangia, such as unite to form the zygospore in Mucor may be a much de- rived form, for there are a number of genera in the Mucorales (e.g., Dicrano- phora and ZygorJiynchus) in which the two uniting gametangia are very unequal in size and appearance, more like the antherid and oogone in some of the Sapro- legniales. Akin to the Mucorales are probably the Entomophthorales and the Zoopagales. In the author's opinion, the Phycomycete line comes to an end there, not proceeding to the higher fungi. The Ascomycetes are believed to have arisen from some algal ancestor re- lated to the Florideae. In this algal group the oogone (carpogone) consists of a swollen basal portion with a receptive trichogyne to which a naked sperma- tium adheres. From the basal portion then grow out hyphae, at whose extremi- ties are produced the carpospores or, in Liagora tctrasporifcra, tetrasporangia. This structure of carpogone, threads, and spores is a spore-fruit wdth, usually, surrounding and protecting vegetative cells. In many of the Ascomycetes occur 2. The terms tinsel flagellum and whiplash flagellum were used by Couch (1941) in the sense that Vlk (1938) used the words Flimmergeissel and Peitschengeissel. They designate respectively the more slender, wavy flagellum with numerous fine lateral threads and the thicker, stiffer flagellum of two parts — a thick basal portion and, at its upper end, a thin lash. These fine details can be observed only by special staining methods or by observation with the dark-field microscope. BESSEY: MYCOLOGY 255 similar oog'ones with triehoiiynes and antlierids i)roduciiiy' nonmotile spermatia. From the fertilized oo servations. What is apparently the same species of small planthopper was described originally from Spartina grass growing on the high dunes of Long Island, and has been taken also on a species of TJniola growing on the high dunes along the North Carolina coast. Here we have, apparently, two different regions with approximately the same physical factors harboring the same species of insect. In Northern Michigan, however, I observed another larger species of planthopper living in tlie sheltered beach pools on rushes, whereas this plant- hopper was not found along the shores of tlie lake where the rushes were sub- ject to high winds. Every student of this order who lias collected extensively in the field has had this experience. Two nitches, wliich are as far as can be judged identical in the more important biological and physical factors, are vastly different in regard to the total population of Homoptera; for the one will yield a large number of specimens whereas the other seems to have none. What then are the factors that make such conspicuous differences? Whether any of these observations will stand the test of carefully planned experiments with METCALF: HOMOPTERA AUCHENORHYNCHA 53] accurate measurements of all of the known factors in the different environ- ments should command the study of future students. That the field of ecology has been too much neglected is abundantly evi- denced. I need point out only a single example. Our studies of the great grassy plains of the Missouri and Mississippi valleys have largely neglected the leaf- hoppers and planthoppers which occur in a normal grasslands area. Yet Os- born's studies showed many years ago that the total population of these insects is of the order of one to two million individuals per acre. Now such an impor- tant observation as that cannot be neglected in studying the sum total of all of the factors, physical and biological, in the environment. There is great need for more careful studies in ecology from all parts of the world. The inference of such studies on the development of the science of ecology and of the economic control of insect pests is incalculable. Careful studies such as are now being made by two Finnish homopterists, Lindberg and Nuorteva, should be initiated by students in all parts of the world. Until about fifty years ago very little attention was given to the economic importance of the Homoptera. However, a few species received some notice; chief attention was given to the spectacular appearance of the seventeen- year and the thirteen-year cicadas and little attention to the conspicuous but relatively inconsequential damage done by the so-called buffalo treehopper. But starting about fifty years ago a sequence of events impressed upon entomologists the importance of the Homoptera in relation to crop damage. One of the earliest and most spectacular of these incidents was the great destruction wrought to the sugarcane fields of Hawaii by the sugarcane planthopper imported from Aus- tralia or New Guinea and its control by introduced parasites. Also relatively early was the damage caused by the sugarcane froghopper in Trinidad. Fol- lowing this was the destruction by the potato leafhopper of potatoes, beans, and peanuts, and the damage caused by the sugarbeet leafhopper to the growing of sugarbeets in the western United States. More recently, the damage caused by the alfalfa froghopper has again emphasized the importance of these insects as pests of agricultural crops. Concurrently with the foregoing, or nearly so, there developed the apprecia- tion of the economic importance of these insects, particularly the apple leaf- hopper complex; various species of cotton leaf hoppers in Africa, India, and Australia; the importance of the grape leafhopper in the United States; of leaf- hoppers on cranberries in New Jersey; and of leaf hoppers and planthoppers on rice, particularly in Japan. Other economic pests perhaps should be mentioned, but most of these are pests of minor crops or are of only local consideration at present. Another development is the importance of these insects as vectors of certain diseases of crop plants. Recent important summaries of these have been pub- lished, and mention should be made of such important diseases as curly-top of sugarbeets and other types of curly-top transmitted by CircuUfer tenellus, of peach yellows by Macropsis trimaculata, of the phloem necrosis of the elm by Scaphoideus, and of various mosaic diseases and several kinds of yellows trans- mitted by leafhoppers. The life histories of many of the economic pests belonging especially to the leafhopper group have been studied but there are many other forms which 532 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES have received only cursory attention. The life history of the seventeen- and thirteen-year cicadas in North America is well known owing to the comprehen- sive studies of Marlatt. Osborn and Ball made very great contributions to the life histories of the leafhoppers in Iowa many years ago, and Osborn studied the life histories of the leafhoppers of Maine and the froghoppers of the same region. More recently some contributions liave been made to the life history of the alfalfa froghopper. Some general studies of the life histories of the tree- hoppers were made many years ago by Funkhouser, and some of the economic pests in this group have been rather generally studied. Much still remains to be done, especially in the tropical regions of the world. The fulgorids have been rather generally neglected; the life histories of only a few species have been studied and these rather incompletely. The phylogeny of the group as a whole is rather poorly understood. Most of our present-day discussions are based upon the studies of Stal, made nearly 100 years ago. Less than 500 of the present known 3,500 genera and perhaps less than 4,500 species of the known 30,000 species were then known. Stal con- ceived the group as comprising four families and for the most part we now con- sider these of superfamily or even higher rank. Basing our studies of the group's phylogeny on such a small area of the total population would be like basing our studies of geography on the knowledge of geographers of the world before the discovery of America by Columbus, or basing our studies of history on only the history known before the beginning of the Christian era. Fairly comprehensive studies of the genera of Fulgoridae by Muir and others, of the Cercopidae by Lallemand, of the jassids by Evans, Oman, and others, and of the Membracidae by Funkhouser give a rather firm basis for com- prehensive study of the phylogeny of these groups. Perhaps what is most needed now is research on the phylogeny of the families and of the groups higher than the families. For the present I believe that the knowledge of the subfamilies, perhaps of the tribes, of most of the groups is fairly comprehensive. What, then, of the future? AVhat the future holds for the field of taxonomy is anybody's guess. Whether other characters will influence the taxonomy of the group as profoundly as the discovery of the impoi'tance of male genitalia has influenced it in the last quarter century remains unknown. Yet I believe that other characters quite as important as the morphology of the male genitalia will be discovered in the not too far distant future. The present tendency is to confine taxonomic studies to a single genus re- stricted to a limited area of the world's surface. Perhaps this is the best method for making progress. It is unfortunate, however, that so few students are suf- ficiently interested in the suborder as a whole to devote their time and attention to the groups higher than genera. Very little progress in taxonomy is going to be made until we have a thorough restudy of at least the external morphology of these interesting insects, correlated perhaps with a study of the internal mor- phology, of physiology, embryology, ecology, and zoogeography. This, indeed, sounds like a comprehensive program but as long as our knowledge of taxonomy is based upon the phylogenetic concepts of Stal of one hundred years ago and as long as we confine the insects of this group to four or five families, just so long will our taxonomic concepts be inadequate, for the consideration not only of the species already described, but of the genera and species not yet described. METCALF: HOMOPTERA AUCHENORHYNCHA 533 We hear on all sides complaints about the rapidly changing nomenclature, and the International Commission is engaged apparently in an attempt to sta- bilize our nomenclature by decrees fixing certain names. How futile this is can be appreciated from a number of apparent facts. First, it is doubtful whether we know more than a third of the genera and species of the Homoptera Aucheno- rhyncha now living in the world. Second, until recently we have had no com- prehensive bibliography of this group. Third, only about a fifth of the families have been covered with an up-to-date catalogue of the genera and species. It might be remarked in passing that although I spend a considerable portion of my time on the current literature, I can just barely keep pace with it. Yet I am foolhardy enough to believe that any attempt to fix names is going to fail utterly; first, because there are not enough workers to study all of the literature of the past and to fix names with accuracy, and second, because the names that are fixed are bound to change with our increased knowledge of the real taxonomy of the group. The changes in nomenclature in systematic zoology are no more drastic than the changes in the nomenclature of any other science, biological or physical, which is developing rapidly. There is something amusing, if not ridiculous, in reading biological papers and noting how carefully the writer has checked every factor involved except the accepted nomenclature of the day. If evolution is an explanation of the facts of the biological world, then the center of origin theory must be accepted. That is, there must be for each species and each genus a center on the earth's surface where these units of the animal kingdom have arisen. It follows, therefore, that a clear understanding of the zoogeography of the animals of a group is a necessary prerequisite to an under- standing of the taxonomy, ecology, phylogeny, and other areas of the field of biology. A great deal of progress has been made in the study of the zoogeography of the Homoptera. Of course, much more than has already been discovered awaits the inquiring mind of the future student. Most of the facts of zoogeog- raphy are so patent that they would seem to need little argument for their sup- port. Except where nature has been interfered with by man and his commerce, we would naturally expect that species would spread from their center of origin gradually, perhaps more rapidly than we think, to other areas to which they can adapt themselves. A firm foundation for our study of zoogeography was established by many different workers working on local lists of the countries of Europe, the states in the United States, South Africa, India, Japan, Australia, various countries in South America, and other regions. The real purpose of a short history such as this is to call attention to the great areas of study which await the nimble fingers and keen minds of future research workers. For these alone can develop the techniques which will push forward the frontiers of our knowledge of one of the larger orders of the insect kingdom and one which contains some of the most bizarre animals known to man. 534 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES HEMIPTERA Egbert L. Usinger University of California, Berkeley The status of our knowledge of the Hemiptera in 1853 can be summarized by citing the best general work of that period, Amyot and Serville, Histoire na- turelle des insectes, Hemiptcrcs (1843). This classical work was built upon the solid foundations of Linnaeus (1758, 1763), Fabricius, especially the Sy sterna Rhyngotorum (1803), Latrielle (1796-1810), Duf our (1833), Burmeister (1835), and Westwood (1840). In the United States, Thomas Say (1831-1832) should also be mentioned with this group of pioneers. In 1853 no general catalogue ex- isted, but at the end of the decade Anton Dohrn (1859) published his Catalogus Hemipterorum. This was the first world catalogue of this great order. Since 1853 tremendous progress has been made. Perhaps the greatest single step was the work of the Swedish father of hemipterology, Carl Stal. Stal was the greatest hemipterist of all time and managed to crowd into his brief forty- five years of life the publication of fundamental works in the Orthoptera, Chry- somelidae, and Hemiptera. Stal had a remarkable sense of fundamental charac- ters and his keys to the principal groups of Hemiptera are still the best keys we have in certain groups. His epoch-making Enumeratio Hemipterorum ( 1870- 1876) in five parts has been called the hemipterist 's bible. Unfortunately this great work did not include a treatment of the difficult and very large family Miridae. Following upon the work of Stal another Scandinavian, 0. M. Renter, de- veloped a classification of the Miridae which filled the great gap in Stal's work. Eeuter devoted himself in the later years of his life to a fundamental phylo- genetic study of the Hemiptera (1910, 1912). China and Meyers (1929) con- tributed further to our knowledge of phylogeny and still later (1933) China gave the latest phylogenetic diagram. Greatly augmenting our knowledge of the Hemiptera of foreign places, the great faunal works appeared during the last half of the nineteenth century. Among these may be mentioned the Fauna of British India by Distant and the Biologia Centr alia- Americana and the Fauna Hawaiiensis. These and other great works expanded our knowledge to all parts of the world and gave a breadth to the classification of the Hemiptera that was not present before this time. Cataloguers were very helpful in the latter part of the nineteenth century, and especially to be mentioned is the great Catalogue general des Hemipteres, Heteropteres (1893-1896) by Lethierry and Severin. The Lethierry and Seve- rin Catalogue is still the only world catalogue for most groups of Hemiptera. Unfortunately it did not include the Miridae nor the aquatic Hemiptera. Atkin- son gave us the Catalogue of the Capsidae (Miridae) (1889), and this is still our best catalogue for this important family. Only recently, starting in 1927, was an attempt made to prepare a new General Catalogue of the Hemiptera. This enterprise was a cooperative one with contributions from scientists through- out the world. A number of fascicles appeared over a period of twenty years. The enterprise was abandoned recently by Smith College, but has been revived by Z. P. Metcalf at North Carolina State College. USINCER: HEMIPTERA 535 The growth of collections marks the development of most of the systematic sciences and this is trne in hemipterology as well. The great collections of the present time are the collections of the British jMnseum (Natural History) in Lon- don, the Natnrhistoriska Riksmnsenm in Stockholm, and the great museums in Helsinki, Vienna, Berlin, Paris, Budapest, Leyden, Genoa, and to a lesser extent elsewhere on the European continent. In the United States great collections were developed somewhat later, and among these may be mentioned those of the United States National IVIuseum, the Museum of Comparative Zoology, the American Museum of Natural History, the Academy of Natural Sciences at Philadelphia, the California Academy of Sciences, the Carnegie Museum, the Chicago Natural History Museum, the Museum at Cornell University, the Snow Museum at the University of Kansas, the Museum at the University of Michigan. Progress in the classification of Hemiptera may be marked not only by the traditional taxonomic works but also by great landmarks in improved ap- proaches to the subject. One of these was the pul)lication by Singh Pruthi on male genitalia in the Hemiptera. This work was published in 1925 and provided a new set of data upon which to base classifications. Another new set of charac- ters was discovered by Tullgrcn (1918) and Ekl:)lom (1928). These authors found the maxillary levers to be of significance in the higher classification of the Hemiptera, and also found that the arrangement and position of the tricho- bothria were of significance in the over-all classification. During the period of taxonomic progress, other students were furthering our knowledge of the biology of the Hemiptera. Among these the first was Dufour (1883). Later Hungerford, Hoffman, Miller, Readio, Butler, and Weber con- tributed greatly to this field. The subject of the physiology of insects was also pur- sued during much of the period covered by this century. Dufour (1833) did the first significant work in this field but classical studies awaited the researches of Wigglesworth. Wigglesworth selected as his experimental animal the bug Rhodnius prolixus. RJiodnius, being a blood-sucking insect, was especially well adapted to studies of this kind because it could be reared in the laboratory and fed only once between each instar. Wigglesworth studied the moulting of in- sects and many other details of the physiology of insects. The subject of genetics should be mentioned because bugs were used very early in the development of this science. The Pentatomids, in particular, were used for cytological studies in the early part of the present century. Recent work of this kind is much more far-reaching and concerns the chromosomes of many other families of Hemiptera. It is too early to say what significance this work may have on our final classification, but certainh^ karyology will provide an additional set of taxonomic characters. Economic investigations have always played an important part in ento- mology, but this has come to be more striking during the twentieth century. In the Hemiptera the most important pests are the bedbug, which was studied from earliest times; the chinch bug, which is so injurious to agriculture in the Middle West and was one of the earliest insects of economic importance to be studied in the United States; lygus bugs, which have come to the fore only in the last few years. Among the numerous other pests are the squash bug and the harle- quin cabbage bug. Of great importance in biological control in the 1920's was a bug of the genus Cyrtorhinus. This bug had the remarkable property of sucking the 536 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES eggs of leafhoppers and was therefore introduced into the Hawaiian Islands to control the sugarcane leafhopper. It is now a matter of historical record that Cyrtorhinus mundulus brought the sugarcane leafhopper under control and saved the sugar industry for Hawaii. Another important group of hemipterous insects is the subfamily Triatom- inae. Triatoma bugs are the vectors of the American trypanosomiasis or Cha- gas' disease. This disease of tropical America was discovered in 1909 by Chagas and since that time many investigators have contributed to our knowledge of the disease and its control. The modern period in systematic studies of the Hemiptera in the United States was inaugurated by Uhler in the latter part of the nineteenth century. It was carried on by Van Duzee, Barber, Blatchley, Heidemann, Drake, Knight, Harris, Fracker, McAfee and Malloch, Hungerford, Hussey, Parshley, Torre- Bueno, Sailer, and by a host of others. Elsewhere in the world outstanding work was developed by Horvath, Schouteden, Poppius, Kirkaldy, Bergroth, Handlirsch, Wagner, Lent, Kormilev, De Carlo, Hoberlandt, Blote, Carvalho, Brown, Bruner, Costa Lima, Jaczewski, Lundblad, Mancini, and many others. It is difficult to anticipate trends, but a look into the future may not be out of keeping at this point. At the present time it may be said that most of the regions of the world have been explored fairly adequately, but that our funda- mental classification, the phylogenetic scheme for the Hemiptera, is still not entirely satisfactory. The basic division into Gymnocerata and Cryptocerata, based upon whether or not the antennae are concealed, is quite artificial. There- fore, we need a comprehensive phylogenetic study of the entire order and this will undoubtedly develop out of work that is now in progress in various parts of the world. Second, we need a collation of the regional works that have been pursued by students in various museums in various parts of the world. At the present time it is possible to go from one European museum to another, or from an American museum to a European museum and find type specimens standing under different names in each museum. The fact is that no one has systematically compared these types and established the synonymy which is so necessary before any really comprehensible classification can be established. Finally, we need a modern catalogue of species and keys to the genera of Hemip- tera for the world. Thus it might be said that the analytical part of Hemiptera classification has been fairly well done but that tlie synthetic part — the bringing together of all the information — remains to be done. Therefore it is clear that the next century has a big, and perhaps the most significant, task ahead, namely, to bring all of the scattered information together into a comprehensible whole. NEUROPTERA AND MECOPTERA F. M. Carpenter Harvard University By 1853 the Neuroptera and Mecoptera were being investigated by a number of well known entomologists. F. Brauer had published more than a dozen papers on them, mostly dealing with life histories, and L. Dufour had made important CARPENTER: NEUROPTERA & MECOPTERA 537 contributions to a knowledge of their internal structure. Taxonomic studies were being carried on by H. Burmeister, J. Curtis, J. 0. Westwood, F. Walker, P. Rambur, and H. Hagen. At that time, of course, the order Neuroptera was an ill-defined assemblage of unrelated insects, including mayfl.ies, dragonflies, termites, bark lice, stoneflies, and scorpion flies, in addition to the insects now termed Neuroptera. Just a century ago E. Newman, following a suggestion made earlier by Erichson, proposed a division of the order. One group (Neuroptera) was to include the insects which we now know as Neuroptera, Mecoptera, and Trichoptera; the other (Pseudoneuroptera) was to contain all the other families previously placed in the order. Although further limitation of the order Neu- roptera has since been made, Erichson's and Newman's proposals were signifi- cant in two respects: they emphasized the difference in the metamorphosis of the two groups of insects thus separated and they anticipated the natural or phylogenetic classification of insects which was more generally applied several years later, following publication of the Origin of Species. The order Neuroptera of Erichson and Newman was usually subdivided by contemporary entomologists into four families : Sialina, Hemerobina, Panorpina, and Phryganina. Ordinal separation of the caddis flies and scorpion flies was gradually made in publications by C. Gerstaecker (1863), C. Gegenbaur (1877), F. Brauer (1885), and N. Banks (1892). From 1850 to 1890 there were only three major workers on Neuroptera and Mecoptera. Brauer continued his studies on their life histories and immature stages, dealing chiefly with Austrian species. Hagen published many taxonomic and biological papers, especially on New World species, his Synopsis of the Neuroptera of North America, With a List of South American Species being the most comprehensive treatment of the group which had appeared up to that time (1886). R. MacLachlan, also, made many important contributions to the knowledge of the world fauna, including a revision of Walker's British Museum Catalogue of Neuroptera and a monograph of the British Neuroptera. Since 1890 there have been many more workers on Neuroptera and Mecop- tera. Nathan Bank's published works, beginning in 1892, is the most extensive and on the widest geographical range of material. H. W. van der Weele has also contributed numerous works on species from many parts of the world, his revisional studies (Ascalaphidae and Megaloptera) being especially important. In more recent years D. E. Kimmins has published numerous papers dealing with the faunas of all zoogeographic regions. L. Navas has described a great many species and L. Kriiger numerous genera, both inadequately and on insuf- ficient material. K. J. Morton, Bo Tjeder, J. L. Lacroix, J. A. Lestage, and P. Esben-Petersen have restricted their studies largely to Old World species, though Esben-Petersen's monographic revision of the Mecoptera (1921) covered all spe- cies known at the time. F. J. Killington, whose British Neuroptera (1936-1938) is truly a classic in the literature on this group, has dealt mainly with British species. Similarly F. Klapalek has published studies chiefly on European Neu- roptera and Mecoptera; R. J. Tillyard on the Australian fauna; R. Smith and F. M. Carpenter on the Nearctic members; and Issiki, Miyake, Nakahara, and Okamoto on Asiatic species. Much of the revisional work done in recent years has been based on detailed structure of the terminal abdominal segments. Studies of this kind, involving 538 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES reworking of type material, have cleared up nuicli of the confusion that has at- tended the taxonomy of both Neuroptera and Mecoptera in North America and Europe, but continuation of sucli investigations is still the prime need. Basic and extensive study collections of Neuroptera and Mecoptera are contained in the British jMuseum (Natural History), which includes the Walker and Mac- Lachlan collections (among others), and in the Museiun of Comparative Zoology, Harvard University, which contains the Hagen and Banks material. Other large museums, of course, also have important study collections, but of more recent origin. Another need, just as important, is studies on the life histories and im- mature stages of these two orders of insects. Virtually only the British and cer- tain European species are satisfactorily known. TRICHOPTERA Herbert H. Ross Illinois Natural History Survey, XJrhana If the starting point of this discussion had been set two years earlier, it could honestly have been said that caddis flies in North America were then represented by only a handful of scattered descriptions. But in 1852^ F. M. Walker de- scribed about 60 species from North America, and this was followed in rapid succession by additional descriptive efforts by Hermann Hagen, Kolenati, and the Abbe Provancher, so that by 1880 some 150 species were described from the North American region. Even with the inclusion of Walker's work there was relatively little known about North American Trichoptera in 1852. Only a few species described by Thomas Say had been illustrated in American scientific literature, while the other species were known only by brief, inadequate descriptions. The European fauna, however, was surprisingly well investigated. Especially notable had been the researches and publications of the Swiss worker, Pictet. In 1834 he gave a fine account of the main groups of the European Trichoptera, illustrating not only many pertinent features of the adults and larvae, but also life-history data on most of the large groups. Pictet divided the Trichoptera into about ten genera, and these anticipated in almost uncanny fashion the major groupings which later became established in the order. Contemporaneously with Pictet, two British workers, Curtis and Stephens, made significant contributions to the recognition of caddis-fly genera, and Zetterstedt added considerably to the knowledge of the fauna of northern Europe. Up to this time, however, the generic and specific diagnoses were on a very superficial level, and information was available in usable form for only a few sections of the European fauna. 1. Here and elsewhere in this article, dates refer only to publications, for which the full references may be found either in Bull. 292, New York State Museum, or in Zoological Record. ROSS: TRICHOPTERA 539 The modern pattern of caddis-fly study was initiated suddenly and deci- sively by the English worker, Robert IMcLachlan, in his monumental treatise on the European caddis-fly fauna (1874-1884). McLachlan defined most of the modern families and genera, introduced genitalic structures as the chief basis of specific diagnosis, and gave a comprehensive set of clear descriptions and illustrations for most of the European species and many of the Asiatic ones as well. McLachlan's work served as a stimulus to a group of energetic students of the order who described species from every part of the globe. Nathan Banks (1892 to 1951) was especially active in investigating North American, South American, and Pacific Island forms; A. B. Martynov (1892-1934) described much of the Asiatic and Oriental fauna, with especially valuable papers on the Siberian forms; George Ulmer (1900 to date) not only studied the Oriental, Neotropical, and African faunas but also wrote the Trichoptera volume (1907) of Genera hisectorum, which has been and still is the starting point of all seri- ous world studies in the order; Longinos Navas (1905-1933), probably the most prolific writer, described material from all areas; and Martin E. Mosely (1919- 1948), whose many papers are ably and fully illustrated by D. E. Kimmins, elucidated the Trichoptera of many lands. Soon after the turn of the century the tremendous upsurge of interest in limnological work added its impetus to caddis-fly studies, especially in the inves- tigations of immature stages. In this field outstanding contributions were made in Europe by Thieneman (1903-1926), Siltala (1900-1908), Wesenberg-Lund (1908-1915), and Ulmer, whose Trichoptera volume (1909) of Brauer's Suss- wasserfauna DeutscMands was of great value for diagnosis. In North America similar studies were reported by Vorhies (1905-1913), Lloyd (1915-1921), and Krafka (1915-1926). A great boon to taxonomic work on the adults was the discovery late in the last century of the clearing or eviscerating action of sodium and potassium hydroxide solutions. This treatment is especially effective in studying the geni- talic structures of insects. One of the first champions of this procedure in Tri- choptera studies was Cornelius Betten (1901 to date). Dr. Betten not only instructed many students in the techniques of trichopterology, but gave North America its finest reference book on the order, his TricJwptera of New York State, published in 1934 as Bulletin 292 of the New York State Museum. Many other workers have been attracted to the order in the last few decades, and these have added the results of their work to the total. In North America the more active have been Tj. J. Milne, D. Ch Denning, and the author. In Europe D. E. Kimmins, F. Schmid, and F. C. IT. Fischer are especially active in the group. Looking over the record, we see that our knowledge of the world fauna has increased from the dozen or so species described in Linnaeus' time to the four or five thousand known today. Tlie list of the North American fauna has grown from eight or nine in Say's time to eight or nine hundred descriljed today. Im- mature stages are known for a surprising proportion of tlie genera (70 per cent in North America). Integration of larval and adult cliaracters has aided tre- mendously in clarifying concepts of classification. Trichoptera is a relatively easy order in which to start studies. There are 540 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES synoptic treatments available for the faunas of several large areas — Europe (McLaehlan, 1874-1884), Russia and Siberia (Martynov, 1934), South Africa (Barnard, 1934, 1940), India (Martynov, 1935, 1936; and Mosely, 1933-1949), Sunda Islands and the Philippines (Ulmer, 1930, 1951), eastern and central North America (Betten, 1934; Ross, 1944), and Australia and New Zealand (Mosely and Kimmins, 1952). Many species of other areas are well diagnosed. The excellence of the literature is a real tribute to the high standards of de- scription and illustration set by the pioneer workers in the group. Three areas of future study beckon the student of Trichoptera. First is the recognition of the many species yet unknown, requiring study of accumulated material and additional collecting in poorly known areas. Second is the need for understanding the identification characters and physiological requirements of larvae, so that these may be used as index organisms and possibly as habitat con- ditioners in the control of pollution and in fish management. And third, there is the need to integrate all this on a world basis, so that we may learn more about the evolution and dispersal pattern of the Trichoptera and apply these findings to the solution of some of the many vexing problems confronting the evolutionist and ecologist. LEPIDOPTERA William T. M. Forbes Cornell University Like most things in the fields of philosophy and science, the serious study of the Lepidoptera starts with Aristotle, who used the cabbage butterfly and the native silkworm (probably Saturnia pyri or Pachypasa otus) as examples of metamorphosis. If we may judge by Pliny, his classical followers added little in fact and nothing in method, and the revival of science after a millenium and a half produced quite a little new factual material, but showed little improve- ment in the casual method of presentation used by such workers as Redi and Aldrovandi, Swammerdam and Leeuwenhoek, Mouffet, and even Petiver. Mme. Merian's little book of fifty plates, with a short text, on an equal num- ber of Lepidoptera with their caterpillars and a word on their biology, makes a step forward in the orderly presentation of the group, and this was soon fol- lowed (1679, 1683, 1718) by a hundred more, giving for the first time a unified picture of the order for any region. In the same period (1705) she also opened the tropical Lepidoptera to our view including the larvae, with sixty plates, from Surinam. 1 The next high spot is the sixth edition of Linnaeus' Systema Naturae, in which he tries out his new binomial system on nearly forty Lepidoptera selected 1. I do not cite the exact titles of these two works, for they differ in the German, Dutch, Latin, and French editions; tliey can be found in Hagen's Bibliotheca, Horn and Schlenkling, or Stuldreher-Nienliuis' biography of Mme. Merian. FORBES: LEPIDOPTERA 541 from his Fauna Suecica, as well as in a few other groups of animals. I very much doubt if the experiment looked as important at the time as it turned out to be. Almost contemporary is Lyonet's Traite Anatomique de la Chenille (1760), so thorough a piece of Avork that, when I dissected the muscular system of the tent caterpillar I found this publication more accurate than any later work; and when Williams rediscovered the prothoracic endocrine gland of the cater- pillar, he found it already figured and discussed by Lyonet! Then came the great period of the picture books, adding up to a pretty clear view of the Lepidoptera of the world. Iliibner's great series on the European Lepidoptera approached completeness in the butterflies and larger moths, and gave a well-balanced view of the micros. Also, his Geschichte is the basis of our real knowledge of the European caterpillars, and his beautiful figures have been copied and recopied right up to modern times. For the exotics nothing remotely resembling completeness was then available, but there were good recognizable figures of all the larger forms from every part of the world, chiefly through the work of Cramer (continued by Stoll) and the publications by Hiibner, with Geyer and then Herrich-Schaeffer, and the appearance of many lesser but still important series, continuing into the century of our present interest. Less pre- tentious in appearance than these illustrated series, but far more scientific in purpose and intended completeness was the French Encyclopedie methodique, beginning with a massive introduction in 1789 and treating (1790-1824) every genus and apparently every known species of Lepidoptera from Alucita to Papilio. It then broke down, with descriptions of relatively few species of Phalaena- and the remaining genera; but the butterflies {"PapiUon") occupy a whole volume. The publication by Smith and Boisduval of a number of Ab- bot's drawings, gave the first clear view of the North American caterpillars, and this was supplemented by less pretentious accounts (primarily of economic species) by Peck, Harris, and others. The last major event before 1853 was the publication of Doubleday's Genera of Diurnal Lepidoptera (1846-1852). This work put the classification of the true butterflies on such a solid basis that the major part of it stands to this day. The skippers have needed more drastic revision — and they really still need it. For the century 1853-1953 we may profitably divide our review into fields of study. Taxonomy The field of taxonomy naturally divides into discovery of kinds, cataloguing, and scientific classification. In 1853 several major works, particularly on the moths, were going through the press. Doubleday's Genera of Butterflies had included a complete catalogue; but Walker was working on his "List" of the moths for the British Museum, which included short descriptions as well as bibliography, and is in fact the last review of the world fauna to be completed. Guenee was working on the moths on a rather more generous scale for the Suites a Buff on, but of this only a few fami- lies were completed, essentially the Noctuidae, Geometridae (Phalenites), and Pyralididae. There has been no complete revision of these three families since, 2. Note that in the Encyclopedie, as in every following publication for nearly a cen- tury, Phalaena, if used at all, meant the geometers, not the noctuids. 542 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES though Boisduval's contributions to the same series of the Papilionidae, Pieri- dae, and Sphingidae are superseded. Herrich-Schaeffer's works were coming out in the same period and supplied figures for many species. The three authors referred more or less to each other, and in some cases no ordinary mortal can tell which author should get credit for a given name, or which is the prior name for a particular species. In the field of major classification each author has a place, but Herrich-Schaeffer, with his more orderly tabulations and keys, has had more influence on later work. Guenee was frequently inspired, but his presentation is less clear, and his attempts to use larvae and biology for classifica- tion were often unsuccessful. Walker was too hurried, and most later workers have found it not worth the trouble to dig out the useful elements of his groupings. This was also the moment at which California appears on the map for Lepi- doptera; for Lorquin went out there in the famous year 1849. By 1852 he had turned back from gold mining to entomology and was sending material to Bois- duval in France. He ranged from Oregon and the Apache country to ''Los Angelos en Sonora," and the results appeared chiefly in Boisduval's two Lepidopteres de la Calif ornie and the "Extrait d'une lettre de ]\I. Lorquin sur la faune de la Calif ornie" {B^dl. Entorn. Soc. France, 1856, p. 98). The noctuids, geometers, and pyralids were turned over to Guenee for the Suites a Buffon. The rest of the half-century was a great period for collecting and describ- ing in all parts of this country, till by 1900 we had a pretty good idea of the North American macros. It is not possible even to list the names — in the East there must have been a hundred workers who made real contributions, in the Rocky Mountain area Snow possibly stood above the others, in Texas Belfrage, on the West Coast Hy Edwards. To me, personally, the outstanding figure was F. G. Sanborn, whose collection, much faded by thirty years of exhibition but still intact, was my first introduction to a real collection of moths. The material collected at this time was worked up by a series of persons, many of them more or less specialists. The bible for the butterflies, sphinges, and bombyces for much of the period was Morris' Synopsis of the Lepidoptera of North America, published by the Smithsonian Institution in 1862. It was intended to follow this with studies of the other groups, but only the Geometri- dae (still "Phalaenidae"), by Packard, actually got published, by the United States Geological Survey in 1876. Packard also continued to work on the "Bom- byces," and the Notodontidae, Saturniidae, and Citheroniidae, including most of the caterpillars, were finally published by the National Academy of Sciences in very luxurious form. The rest of the plan, however, disintegrated. The Noc- tuidae eventually fell to Smith when he came to work at the United States Na- tional Museum, and quite a few fragments were published, mostly after the end of the century; while the micros, which were Fernald's portion, were repre- sented by the crambids and pterophorids, published by the State of Massachu- setts, and by a bibliographic catalogue of the Tortricidae, which appeared in the Transactions for 1882. This was the period when the butterflies became a major specialty. In addition to innumerable scattered papers, the principal manuals were by Morris, his Syn- opsis, already mentioned, by French and others, culminating in the great works of W. H. Edwards (1868-1884) and Scudder (1889), with their rich illustration and vast data on early stages, Edwards mainly on the western, Scudder only FORBES: LEPIDOPTERA 543 on the eastern species. Holland's Butterfly Book (1898) started a new era, for it first figured practically all the butterflies, for both East and West, and at a price the public could easily pay. (It also started the present writer on the Lepidop- tera.) In the present half -century local butterfly books have continued to ap- pear. J. H. Comstock's Hoiu to Know the Butterflies (1904) was more complete on early stages and more compact than Holland, but served for the East only. On the West Coast Wright (1905) would probably have replaced Holland, if the earthquake of 1906 had not destroyed most of the edition. It was perhaps less critical than Holland, but more richly illustrated, and is our chief record of identifications current on the Coast before the fire. For instance, his figures of Pamphila ruricola, and J. A. Comstock's figures can be reconciled with the original description, whereas the supposed type and more recent identifications (e.g., of the brown Atrytone vest^'is) cannot. In the most recent period J. A. Comstock's Butterflies of California and Klots' Field Guide to the Butterflies will probably dominate their respective areas. In more scientific classification rather than the discovery and identification of species, another series of authors and works have dominated the field. Here two works stand above the others, even from a world point of view: Doubleday's Genera for the first clear picture of the world classification as a whole, and Scudder's Butterflies of Eastern North America for the only integration of characters of all stages. Except in the skippers and special studies of limited groups the only other work worth mentioning is Sehatz's Exotische Schmetter- linge: Familien und Gattunge7i der Tag falter (1892), which is roughly the generic part of Doubleday, revised, extended, and brought up to its date. At that time the genitalic and larval characters had not been properly studied for the definition of genera and higher groups, and the time is now more than ripe for another Doubleday or Schatz. Schatz died in the midst of his work, and the classification of the Lycaenidae by Rober represents a lower level of quality. The major classification of the skippers has had a separate history. Doubleday did little with them, Schatz and Rober omitted them, and their serious study prac- tically begins with Scudder. Druce, in the Biologia (1893-1901), and AVatson (1893) extended the scientific approach to a world point of view. More recently Lindsey, Bell, and Williams have given us an integrated picture for the United States (Denison Univ. Bull. Journ. Sci. Labor., Vol. 26, 1931). But Evans' World Revision will be the definitive work : the Africans were published in 1937, Eurasians and Australians in 1949, Americans are beginning to appear, and we hope the rest is in press. All the critical work on skippers has included the genitalia, starting with Scudder and Burgess in 1870; but knowledge of early stages has been too fragmentary for any one to go much farther than Scudder did. Outside the United States the chief region where the butterflies are a special study has been England, I suppose because only in English are there distinc- tive words for "moth" and "butterfly." Here the works are far beyond count- ing; I might only mention that I turn most often to South 's Butterflies of the British Isles. Classification below the species has gone farthest also in the butterfiies. Here we have had a rather violent change in point of view. In the first half of our century most workers who went below the species were chiefly interested in bio- 544 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES logically significant or striking variations, and most of the proposed varietal names represented seasonal or dimorphic forms, or even pure aberrations, if these were striking enough to make a good show in a collection. This was still the almost universal practice when Holland's Butterfly Book was published (1898) and is strikingly evident in the many studies of infraspecific variation by Edwards. But long before, some collectors who had material from widely separated localities, noticed the fact of local variation and began to view locality records as something more than a mere convenient reminder of where to go for more of the species. Chief among these collectors was Staudinger of Dresden, who was comparing material from Mediterranean and northern Europe or blocks of material from central Asia with both; even in 1861 (the Staudinger and Wocke Catalogue) he was practically restricting the term "varietas" to such localisms. In his 1901 catalogue (Staudinger and Eebel), this was done as consistently as possible, and other types of variation were reduced to the designa- tion "ab." Jordan, in England, adopted this definition, and it became rather general in Europe long before it was taken seriously over here, so that when the rather ambiguous terms of the International Code appeared, the official interpretation of the term "subspecies" soon became practically the "varietas" of Staudinger. AVorkers on the American fauna, even German workers on South American material, found the distinction impractical and never adopted it fully, though some tried to take advantage of the rules by calling the old traditional varieties "subspecies," especially where, as usually in mimetic South American types, there was a certain tendency to local restriction. This shows most strikingly in Stichel's revision of the heliconian butterflies in Das Tierreich (fasc. 22) ; but must be considered even in interpreting the let- tered forms under the numbered species in McDunnough's Check List. The au- thor has found a curious complication in Junonia, where racial limits appear to be somewhat different in the two biological phases of the buck-eye. As a result, before 1900 most of the infraspecific work was oriented to seasonal or genetic variation or to direct response to conditions, whereas most recent work has been on local variation, which can be more easily equated with the code concept of "subspecies." The most intensive studies have been a long series of papers by Eoger Verity on European butterflies, largely aimed at tracing the presumable lines of migration of populations in past ages, and in America such studies as those of Gunder and of Hovanitz on local variation in California species of Melitaea and the species and near-species of Colias on both continents. Verity's great works are the Rliopalocera Palaearctica : Papilionidae et Pieridae (1905- 1911); Le Farfalle diurne d'ltalia: Hesperides in 1940; and his study of the Lycaenidae in 1943, with an amazing series of colored figures. But for the geog- raphy of the Nymphalidae we must still refer to his scattered papers. He belongs to the school which analyzes local variation on three levels : the race proper (which he calls exerge), the subrace (his race), and of course the unnamable field form. sphingidae: The history of the Sphingidae for the century is short and simple, and for the most part distinct from that of any other group. When the century began, the authority for the United States was Harris' monograph (1839), cited above; then came Morris in 1862, and Boisduval's world revision in the Suites a Buffon (1874). In 1886 Grote and Fernald both published reviews based on Boisduval; Grote's was the one that covered North America, but my own early guide was Fernald's Sphingidae of New England. FORBES: LEPIDOPTERA 545 The Rothschild and Jordan revision, published as a supplement to the Novitates Zoologicae in 1903, was a turning point, for it first put the classification on a solid basis, with keys as well as short descriptions for the whole world, and proper consideration of the geni- talia. However, his names were applied according to an odd code of his own; when his use of an older name agreed with either tradition or later rules, it was pure coincidence. Early stages were also practically neglected and have never yet been studied from the world point of view. I studied the larval characters in 1911 (Ann. Entom. Soc. Amer., 4:261-280) under the encouragement of Beutenmuller, who had got together a good many specimens for a study of his own, then abandoned. Later I saw a few of them again in a most unexpected place. The only scientific approach to the pupae is by Mosher (Ann. Entom. Soc. Amer., 11:403-442, 1918). The beautiful and detailed figures of Moss and of Bell and Scott ("Sphingidae of Peru," Trans. Zool. Soc. London, 20:65-118, 1912; Nov. Zool, 27:333-424, 1920; Fauna of British India, "Moths," Vol. 5, 1937) give us a rich but superficial view of the exotic fauna. Since, Beutenmiiller's work on the adults has been mainly a modification of Rothschild and Jordan; but B. P. Clark's series of papers in the Proceedings of the New England Zoo- logical Cluh have some important data on races in the United States, and have added a few, but very striking, species to our knowledge of other parts of the world. At the moment we have a fuller and sounder knowledge of the Sphingidae than of any other family of moths, yet scientifically the early stages are almost a blank, there being only those two hurried papers mentioned above. Miss Edna Mosher's on the pupae and mine on the larvae, each limited to a partial sample of the Holarctic fauna. saturxioidea: Next to the sphinges, the saturnids are probably the most popular group of moths, but their discussion more properly belongs under biology rather than taxonomy, for knowledge of their early stages and biology has always anticipated their classification. In 1853 I suppose most people in the East knew them through Harris' Insects Injurious to Vegetation; and Boisduval supplied two California species from Lorquin's collecting. Clemens' revision in Morris' Synoi)sis then became the authority, and the four genera recognized by them (Saturnia and Attacus, Ceratocampa and Dryo- campa) were the names familiar to amateurs until Holland's Moth Book came out in 1903. In fact, the saturnids were almost a specialty for amateurs and dealers, who knew how to find the cocoons, and who published some of the life histories in great detail. My own authorities "before Holland" were Harris' Insects Injurious, Mrs. Ballard's Among the Moths and Butterflies (1890), Mary Dickerson's Moths and Butterflies (1901) and Eliot and Soule's Caterpillars and Their Moths (1902). What Westerners did, I have no idea. But when Holland came out, we had colored figures of everything for the country, though we still used the four amateur authorities for biological data. On the scientific side, the classification has never come properly into focus. Packard's revision for the National Academy was unfinished when he died. In its final publication it was rich on early stages, but fragmentary in classification. In Europe the picture was similar: plenty of material in the hands of dealers, including early stages, plenty of figures, and very little classification. Only this year have we at last a world classification (Michener, "The Saturniidae (Lepidoptera) of the Western Hemisphere," Amer. Mus. Nat. Hist. Bull.. Vol. 98, art. 5), which actually ties in most of the Eurasian types and leaves only the Africans incomplete. But still little has been done to work up the rich and significant larval and pupal characters. One might add that, in general, the family limits have been clearly understood for Europe and North America; South America, however, seems to have been a problem for many earlier authors, including Kirby in the Catalogue of Lepidoptera Heterocera (1892) who included most of the relatives of the lo moth in the Lasiocampidae, along with members of several other very distinct families. BOMBYCEs: That array of unrelated but similar families known colloquially as the bombyces have had too complex a history to follow in detail. In America the authority, as the century opened, was Morris; in Europe the second volume of Herrich-Schaeffer (1845) was available, and this was soon supplemented by Heinemann's Schmetterlinge Deutschlands und der Schiveiz (1859), but during the whole period in Europe picture books have dominated the more serious classifications. In America the publication of 546 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Neumoegen and Dyar's series of papers in the first two volumes of the Journal of the New York Entomological Society was an important event and so, in a less formal way, was Stretch's Zygaenidae and Bombycidae, published in parts. This last remained far from complete; the plates alone were finally published without color in the Journal of the New York Entomological Society in 1906. Hampson's Catalogue of the Lepidoptera Pha- laenae (1898, 1900, 1901, 1914, 1920) gave us a world view for a few families (Euchro- miidae, Nolidae, Arctiidae, Agaristidae, s. str.), but for the rest we still have only the colored figures and short descriptions in Seitz. In the first part of our century, and in earlier periods, this group was believed to be natural, being the Phalaena Bombyx, with a little of the Sphinx, of Linnaeus; but it was gradually realized to be heterogeneous, and the history of its major classification is that of the order as a whole. noctuidae: The Noctuidae start our century in wonderful confusion, which has not yet been wholly cleared up; for Guenee and Walker published world revisions, while Herrich-Schaeffer, followed by Lederer, studied the European types more fully and pre- sented keys. Each author divided the group into a series of families, but none defined them clearly, and no two wholly agreed. Also, it was already recognized that the deltoids belonged with the Noctuidae, rather than with the pyralids, but only Herrich-Schaeffer and Lederer made the union definite. There were also wide divergences in the use of generic names, which were refiected in this country by the divergent usages of Grote and Smith, followed later by Dyar and Hampson. Most works up to the First World War followed tradition more than rules, and diverged in their use of both ; finally, after the war, more and more authors began using the code of 1911, but their individual interpretations added to the general confusion, and the rules often resulted in still further divergent uses of the older names. At the moment, from the world point of view, we have about three-quarters of the family in Hampson"s Catalogue of the Lepidoptera Phalaenae and in Seitz, a complete view of the Palearctic fauna (such as it is) in Seitz, and the rest in fragments: the North American deltoids by Smith (1895); a catalogue including also the South Americans by Schaus (1916, with a key to genera) ; the fauna of British India (1895) ; and a host of loose descriptions. In the pseudodeltoids we actually have nothing since Walker — which means nothing at all, for very few were known then. The subdivision of the family falls rather sharply into two periods. The early work- ers, like Guenee and Walker, divide it into a large number of weakly defined families; Lederer (1857) already saw it as a unit, but takes up these "families" in discussion; and through the rest of the nineteenth century we have general recognition of the family as single, but a similar protean series of groups, mostly treated as subfamilies. Hampson (1903) presented a new system of subfamilies, based chiefiy on certain points used as key characters by Lederer; and these, though recognized as partly artificial, have been con- venient enough to serve up to the present. The Noctuidae even more than the skippers have been a main line in the study of genitalia. In 1857 Lederer was already examining all the available species and figuring the tips of valves. Smith, who for some decades was best known in this country for their study, also limited himself as a rule to the valves, usually pulling out and mounting a single valve, when he intended to save the specimen. About 1909 both Smith (with Grossbeck as technician) and Pierce in England began making more complete dissec- tions on a large scale, and the younger group of workers have brought the technique to a very high level. (I was in the chain; Grossbeck taught me in 1910, and I showed some of the rudiments to Pearsall and Busck.) geometridae: The geometers started the century just like the Noctuidae, with world reviews by Guenee and Walker and more precise analyses of characters by Herrich- Schaeffer and Lederer. Packard (following Guenee's system) gave us our bible for the family in 1876. The present system of subfamilies was established by Meyrick in 1892 (Trans. Entom. Soc. London), and adapted to our fauna by Hulst (Trans. Amer. Entom. Soc, 23:245-386, 1896); and except for some primitive oddities can be considered fully natural, not a merely convenient grouping like the Hampson system in the Noctuidae. More recent work is scattered, and pretty tentative as regards tribes and genera; it takes the form of many small papers, and the fraction of a world revision written by Prout and published by Seitz. Work on genitalic characters is fragmentary and largely unpub- FORBES: LEPIDOPTERA 547 lished; that on early stages is mostlj^ superficial, though studj^ of the pupae is beginning to show some more significant characters. PYRALiDiDAi:: ThesB again start with the same pattern: Gueuee and Walker, Herrich- Schaeffer and Lederer, with Guenee introducing the system most used during the nine- teenth century, and Lederer foreshadowing the system used most recently. But in these Pyralids Lederer did not finish his work, covering only the subfamilies grouping about the Pyralidinae and Pyraustinae and omitting the crambid and phycid-like types. This time the modern pattern of subfamilies and genera goes back chiefly to Hampson, in a series of catalogues (1895-1899), merely listing the species in most subfamilies, but describing and figuring the species of Phycitinae, Anerastiinae, and Galleriinae, with Ragonot, in volumes 7 and 8 of the Romanoff Mevwires. Beginning with this group Seitz fails us completely; and for species outside the last three subfamilies we have nothing beyond Guenee and Walker except the little group of revisions for North America follow^ing the break-up of the plan for a North American monograph: the Pterophoridae and Crambinae by Fernald, the Nymphulinae and Scoparias by Dyar, the few Macrothecinae by McDunnough. microlepidoptera: The micros have followed a very different pattern, and a more complex one. American zoologists for the first fifty years usually treated the smaller species almost wholly from the point of view of biology, and there was a strong feeling that, in the larger genera like Coleophora, Lithocolletis, and Nepticula, adult characters hardly existed. Meanwhile a few stray workers were considering and describing the adults, but only three of these had any real Infiuence: Brackenridge Clemens, especially after Stainton had reprinted his work as the Tineina of North America (1872), V. T. Chambers a little later, and Lord Walsingham, with his series of papers resulting from his trip to California and Oregon in the early 'eighties. When I started, the conventional way to "determine" a tineoid was to rear it, look up the food in Chambers' catalogue {Bull. U. 8. Geol. Snrv., Vol. 4, no. 1, art. 4, 1878), check with Stainton's Natural History of the Tineina for the genus with similar behavior in Europe, and then come up with a guess at the species — the guess was occasionally correct. If it was a broad-winged species with less distinctive habits, we would cruise through Clemens. In Europe the micros were arranged in orderly fashion somewhat earlier. In 1853 Herrich-Schaeffer completed the Lepidoptera, with keys and many figures, as a supple- ment to Hiibner's Europeans. Somewhat later Heinemann reworked the fauna of central Europe (finished in 1877); and the picture books figured enough species to be service- able. There was also the series of volumes of Stainton's Natural History of the Tineina, with their great contribution to the biology. Then, in the 'eighties and 'nineties Meyrick, in working out the Australasian fauna, developed an ai'bitrary but useful scheme of families for the micros, which he applied to the European fauna in his Handbook of 1895; and this was adapted to the American fauna by Busck in Dyar's List. Meanwhile Spuler had been working in Germany on a more natural system for the micros, partly in col- laboration with Comstock's work on the macros; and the result appeared in Hormuzaki's Analytische Uebersicht der palaearctischen Lepidopterenfamilien (1904) and more fully though without any keys, in Spuler's own Schmetterlinge Europas (1910). We adapted it to the American fauna in the Manual for the Study of Insects, which then became again An l7itroduction to Entomology, and the first part of the Lepidoptera of New York (1920, 1924). The Introduction has in fact the later version, since the Lepidoptera was about four years in press. Scattered recent studies show the time is ripe for another reworking. In the last half-century there have been a number of helpful revisions and catalogues, mostly of single families and altogether covering hardly half the micros. For central Europe we have Hering's contribution of the Lepidoptera to Brohmer's Tierwelt Mittel- europas ; and for the whole world we have Fletcher's catalogue of all the genera, with their references, types, type localities, and all their synonyms; also their families according to the Meyrick formula. The following list gives a summary of what we have. Note that the Lepidopterorum Catalogus (Lep. Cat.) is supposed to have a complete bibli- ography and general localities, but no descriptive matter; the scope of the Genera Insec- torum {Gen. Ins.) is also world-wide, and gives descriptions and keys down to genus, but, as a rule, only original references. All but the Stenomidae are by Meyrick. The other works cited are for the Nearctic region only. 548 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES TOBTRiciDAE (s. str.) Lep. Cat., 10, 1912; Gen. Ins., 149, 1913. OLETHEEUTIDAE Heinrich, U. S. Nat. Mus. Bulls. 123, 132, 1923, 1926. PHALONiiDAE Busck, Joum. N. Y. Entom. Soc, 15:19, 1907 (omitting species of Phalonia). CARPOsiNiDAE Lep. Cat., 13, 1913; Gen. Ins., 179, 1923. ypoNOMEUTiDAE (and PlutelUdae) Lep. Cat., 19, 1914. GLYPHiPTERYGiDAE Lep. Cat., 13, 1913; Gen. Ins., 164, 1914. HELiODiNiDAE Lep. Cat., 13, 1913; Gen. Ins., 165, 1914. GELECHiiDAE Busck, Proc. U. S. Nat. Mus., 25, 770-930, 1903 (to species) ; Meyrick, Gen. Ins., 184, 1925. CECOPHORiDAE Busck, Proc. U. 8. Nat. Mus., 35, 189, 1909 (to genus only); Meyrick, Gen. Ins., 180, 1923. BLASTOBAsiDAE Dietz, Traus. Amer. Entom. Soc, 27, 100 ff ; 1910. STENOMiDAE Busck, Lep. Cat., 67, 1935. COLEOPHORIDAE (NE. U. S.) Heini'ich, in Lep. N. Y., 202-217, 1924. GRACiLARiiDAE Lep. Cat., 6, 1912; Gen. Ins., 128, 1912; Ely, Proc. Entom. Soc. Wash., 19:29-77, 1917 (U. S. genera and catalogue of species). TiNEiDAE Dietz, Trans. Amer. Entom. Soc, 31:1-96, 1905. ADELiDAE (long-horns only) Lep. Cat., 6, 1912; Gen. Ins., 133, 1912. MiCROPTERYGiDAE (and Eriocrauiidae) .. Lep. Cat., 6, 1912; Gen. Ins., 132, 1912. Major Classification The major classification and pliylogeny of the order together have had rather a separate liistory. At the beginning of the century ideas of evolution had not be- come general, and most people were satisfied with approximations to the Linnaean system, supplemented by suggested cross-resemblances between the various groups, such as are represented in the diagrams in Herrich-Schaeffer by a web of lines and circles (e.g., vol. 6, pis. 1, 7, 15). Even in lierrich-Schaeffer's time it was realized that the "Bombyces" and "Tineina" were congeries of perhaps unre- lated forms; yet the groupings are such a convenience that they are used even now to some extent. After the Darwinian theory was digested, weblike classifications were recog- nized as artificial and there was a serious search for characters marking primi- tive or specialized forms, and indicating the lines of development. The most important early American work was by Packard, most fully published in the introduction to the Monograph of the Bomhycine Moths (1895). In the same year Comstock published the Manual for the Study of Insects, with a key to the families defined on modern lines, and also a phylogenetic arrangement, notably breaking up the bombycine families and distributing them according to their true relationships. In the same period in Europe Spuler (from 1892) was work- ing on adult, and Chapman (from 1893) largely on pupal, characters. Dyar came along immediately afterward with a more complete study of the larvae in a series of papers, starting with his "Classification of Lepidopterous Larvae" in 1894. The most productive point in Packard's study was the recognition of the very deep character of the differences between a few primitive families, in contrast to most of the order. Comstock became best known for his emphasis on the marked change of structure of the hind wing which set off the earlier "Jugatae" from higher types; but his distinction of "frenulum-losers" and FORBES: LEPIDOPTERA 549 "frenulum-conservers" has also turned out to be significant, when checked to the egg and larval characters discovered or emphasized by Dyar. It was only neces- sary to realize that the frenulum-losers included not only the families that had usually lost the frenulum, but also those, like the Geometridae, that showed merely a tendency to lose it (as worked out in Comstock's A71 Introduction to Entomology [1920] and my own Lepidoptera of New York). Tutt, in his British Lepidoptera, presented a second system, especially in pages 102-128, of volume 1. His idea was that the Lepidoptera followed not one but two roughly parallel lines of evolution from the lowest to the highest branches. His wealth of argument and data gave him considerable authority for a time, but I think he no longer has any followers. Later study of the auditory (or sonar) organs gave further emphasis to the "frenulum-conservers" as a unitary group. The organ itself has long been known, was first described in detail by Swinton in 1877, for a noctuid (Entom. Monthly Mag., 14:123), and I got a glimpse of its phylogenetic value in 1916 {Psyche, 23:183), but it was not until Eggers' studies (1919, 1925, 1928) that its value was established. The organ is also useful below the family level, as has been shown by Richards {Entom. Amer., vol. 13 (no. 1), 1933), working on the Noctuidae, and by Luh (thesis, published only in abstract) on the Arctiidae. At present Kiriakoff is working on other Noctuoidea. Personally I stand by the system of the Lepidoptera of New York and the Encyclopaedia Britannica except for the micros, where recent work (notably that of Hinton) will probably result in some radical changes. But I fear no con- ventional classification will fully express the step-wise evolution of the forms ly- ing below the Tineidae. Some figures : The following counts will suggest the gradual increase in knowl- edge of the species of Lepidoptera. It is rather curious that the last complete catalogue of the order (Walker) lies in just the same period as the first for North America (Morris). All figures are rough, and the suggested totals for the world are merely guesses. The Harris catalogue (1833) was for Massachu- setts only, and the total should be increased to allow for the Abbot discoveries published by Hiibner and Smith; tlie Morris catalogue included Mexico and the West Indies. WORLD FAUNA Noctuidae Linnaeus ( 1758 ) 66 Fabricius (1793-1794) 380 Hubner (about 1820) 784 Walker (about 1860, roughly) 5,625 Hampson (about 1910, partly estimate) 14,357 Present (pure guess) 18,500 25,000 80,000 NEARCTIC FAUNA Harris, 1833 (Massachusetts only) 107 Morris, 1860 (North America) 486 Grote, Edwards, Chambers (about 1880) 1,409 Dyar (1903) 2,128 Barnes and McDunnough (1917) 2,532 McDunnough (1938-1939 ) 2,693 Micros Total 104 535 521 2,817 709 4,198 8,800 33,600 51 428 236 1,551 1,482 4,544 2,346 6,622 3,439 8,495 4,369 9,876 550 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES It appears that the Noctuidae (also the butterflies) are approaching com- plete discovery in the Nearctic, but that the other groups are due for substantial increase. The European list indicates that eventually there will probably be more micros than macros. Morphology The development of knowledge of the external morphology of the Lepidop- tera has been almost wholly of two kinds, either incidental to general studies of the insects, like Crampton's work on a couple of types of Lepidoptera in 1908 and many later papers, or else by-products of classification studies. The study of internal anatomy, however, has been independent. For the caterpillar there has been nothing during the whole century in the class of Lyonet's work in 1760, and for the adult, the dissection of the Monarch, published by Scudder in the Butterflies of Eastern North America (pi. 62, 1888) stands alone. Further work on the anatomy has been voluminous but widely scattered; the fullest and most recent summary is that by Zerny and Beier in Kukenthal's Handbuch (vol. 3, pt. 2, 1936). It shows a fairly complete knowledge of the anatomy as a type, but there is still little on variation of structure within the order. Physiology Physiologists as a rule make slight distinction as to the form they use, whether Neurospora or Paramecium, Brosophila or man; only occasionally has a lepi- dopteran been chosen as an object, and I think never for the sake of contrast with other organisms. Quite recently Carroll Williams has been using cater- pillars, chiefly the cecropia in the study of hormones and their relation to transformation or the mechanism of respiration and hibernation, with their controlling enzymes. Work on the nature of coloration has been more concen- trated on the Lepidoptera, and it has been carried on for a longer period. Ma- son pretty well settled the problem of structural colors in 1927, followed by an actual electron photograph of the structures by Richards. But the question of pigments has been much more complex, though in recent years a number of workers, chiefly English, have done a good deal. The matter of pattern, as distinct from color, should probably be considered morphology rather than physiology, since, though the elements generally appear in pigment, the pattern has the same fixity as morphological characters; it evolves from group to group in a similar way and is frequently foreshadowed by small but definite differences of structure in the individual scales. The realization that pattern elements have a fixity higher than the species or genus came pretty early. In America we are apt to associate it with Smith's diagram for the Agro- tids {Bull. U.S. Nat. Mus., vol. 38, 1890), but at the very beginning of our cen- tury Guenee had a labeled type pattern {S2)ecies General, Noctuelites, pi. 1, 1851). The names of elements have been regularly applied to similar lines and spots in other families, but it has only been gradually apparent that many of these elements are homologous over a wide range of families. For the butterflies in particular an independent nomenclature has been developd, most fully worked out by Schwanwitsch (numerous papers, but the one on the Catagranuna group [Trans. Zool. Soc, 1939, pt. 2] is best known). It is for the future to FORBES: LEPIDOPTERA 551 show whether the nymphalid and noetuid schemes can be homologized definitely, but the absence of similarly definite schemes in the skippers, Castniidae, and Cossidae reduces our hope. The little work done on the physiological forces be- hind the forming of these patterns is too scattered to summarize. The knowledge that coloration and patterns were protective by matching normal backgrounds, greatly antedates our time, but mimicry was a discovery of nearly one hundred years ago and was impressed on Bates (1862) by the wealth of examples he saw along the Amazon. A few years later, Miiller was also impressed by the many cases he saw, farther south in Brazil, in which more than one member of a pattern appeared about equally protected, and proposed what we now call IMullerian mimicry (convergence of pattern to simplify the learning process, and thus to reduce the number of individuals sacrificed by in- experienced predators). Realization by North Americans that mimicry is also (though feebly) a North American matter dates from Scudder's Essay of 1889. Looking back we can date mimicry in a negative way to Linnaeus, for undoubt- edly it was the handling of unrelated models and mimics with similar patterns that caused him to abandon, in his tenth edition, the very useful distinction of four-footed and six-footed butterflies. For a full analysis of the problems involved in coloration, the critical work is certainly Gerald Thayer's Concealing Coloration, and the date is 1909. This work showed fully the functions of concealing pattern and color, mimicry, both tentative and developed, and added flash colors, ruptive pattern, and counter- shading — all largely illustrated by the Lepidoptera.^ Later work has added many details, but little theory, much of which is summarized (without deep insight) in volume 2 of Schroeder's Handhuch (1929). In the field of genetics the Lepidoptera have served from time to time, mainly at first in the breeding of families of specimens to obtain lots with aber- rant patterns, and lots of material distributed by several dealers in the period after 1900 have been better known than the widely scattered publications. Sei- fert did some significant work in the early 1900's showing Mendelian inheritance, but his published reports in 1901 and 1905 do not deal with the genetics, which must be studied from his material preserved at the American Museum. AVhiting worked on Ephestia kiiehniella, and published some data on the genes in 1919, but soon abandoned the moth for its hymenopterous parasites. But the most important work based on the Lepidoptera was that on the gypsy moth, carried on by Goldschmidt over a period of many years, which threw light on the physi- ology of variation and the control of sex. It is summarized, with much other related material, in his Physiological Genetics (1938). Geography The Lepidoptera are a very important source of data for zoogeography, since in various groups we understand the classification well enough to distinguish between true relationship and parallelism; the material is widely collected, and the means of distribution are pretty well understood. Also, from the days of Wallace and Bates we have had workers interested in both Lepidoptera and 3. An interesting side note is the fact that Theodore Roosevelt used his term "nature- faker" chiefly of the Thayers — and it was they who turned out to be right. 552 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES geography, whose views were partly based on what they saw of the butterflies. But much work has been invalidated by being based on false ideas of rela- tionship. The most pretentious publication, Pagenstecher's Geographische Ver- breitung der Schmetterlinge (1909) must be used with great caution, since at that time it was not yet possible in many cases to distinguish between relation- ship and parallelism, and current classifications were arbitrary systems for con- venience in many instances in which he thought true relationship was intended. Schroeder's Handhuch (1929) also has a long chapter on zoogeography, based to a great extent on the Lepidoptera, but here again species lists are often pre- sented without understanding. On a smaller scale we have studies of the spread of an immigrant in a new territory, such as Scudder's work on Pieris rapae {Butterjiies of Eastern North America, p. 1175, Mem. Boston Soc. Nat. Hist., 4:1), and the more recent governmental studies of the spread of the gypsy moth and the European corn borer (the last-named work very superficial). In Europe we have more detailed studies, based on the fuller data available, like Verity's papers on the significance of geographical variation, cited above, and Bryan Beirne's Origin and History of the British Fauna (1952). Biology and Early Stages Our story started with life histories: Aristotle's cabbage worm and silkworm, then Merian's one hundred fifty life histories from central Europe, and a some- what smaller number from Surinam. For the European fauna a large number of naturalists made contributions to the Lepidoptera; but for the mere record of the appearance of the caterpillars, Hiibner's Geschichte marks a high point, not touched before or since. After Hiibner, the steady flow of contributions to life history continued, being integrated by Hofmann and again in Spuler's Schmetterlinge Europas, which shows plain signs of its background — Hiibner, Hofmann, and post-Hofmann. For the more restricted fauna of the British Islands, manuals have come out every decade of the century, but the works of Stainton, Buckler, and Tutt must be mentioned. In America the early work of Abbot in Greorgia was mentioned earlier. Later, about 1900, there was great amateur activity in the northeast, best represented by the three popular works of that time, (Ballard, Eliot and Soule, Dickerson), already mentioned. For structure of early stages we have Fracker on the larva and Miss Mosher on the pupa, published by the LTniversity of Illinois in 1915 and 1916, respectively — studies which are a chief foundation of our modern classi- fication— and also later papers by Miss Mosher on the Sphingidae, Saturnioidea, Notodontidae, and Geometridae. Finally, we have Peterson's Larvae of Insects, of which part 1 (1948) deals chiefiy with the Lepidoptera. In the foreign field data are still more scattered, but for some regions we have unified blocks of material: Matsumura's 6000 Insects (1931) and the Nip- pon Konshu Zukan (1932), Lepidoptera by Esaki and others, for Japan; Bur- meister's Description Physique de la Repuhlique Argentine (1878) or the Lepi- doptera parts of the Fauna of British India (1892-1947, and far from finished), especially the revised volumes on the Papilionidae, Pieridae, Nymphalidae, s.l., except Nymphalinae, and Sphingidae. For biology in the more restricted sense, the literature is so extremely scat- FORBES: LEPIDOPTERA 553 tered that we can only touch on the general works: Packard's Guide for the Study of Insects (1870), Berlese's Gli Insetti (1912-1920); Hering's Biologie der Sclimetterlinge (1926), the second volume of Schroeder's Handbuch (fin- ished in 1929), and Bourgogne's chapter on the Lepidoptera in the Traite de Zoologie (1951). Economic Entomology A review of the history of economic entomology is not a part of this report, but one may note that in this field also the Lepidoptera play a large part. The first serious report was probably Peck's article on the spring cankerworm {Mas- sachusetts Magazine, 1795), which rated a frontispiece. Our century was marked by the inventions of high-pressure spraying apparatus to reach the gypsy cater- pillars in the tall elms of eastern Massachusetts; and the caterpillars still hold their own as test objects, now that economic entomology has gone over from the study of insects to the study of spray chemicals. Looking Ahead This is a very difficult time to look ahead. One can try to extrapolate the present trends, making allowance for those that will last long and those that are ephemeral, or one may remember that our civilization is more than three quarters through the Petrie cycle, and that the next Dark Century is due in less than two hundred years. The first prophecy is that there will be no lack of unknown material to work on. In a few groups, such as the fauna of west and central Europe, or the but- terflies, bombyces, and noctuids of this country, there are few species to add, but in the micros here, and in all groups over most of the earth, even species- making is far from finished. I estimate that we know more than 90 per cent of the micros of Europe, well over half for the United States and Canada, but only a sample (mostly of those that can be easily caught and do not have to be reared) for the rest of the world. For geographical study, the general laws of distribution are known, but they have been applied to the Lepidoptera only in a rudimentary way. It is high time for a new "Butterfly Geography" based on the better known groups, such as the butterflies, sphinxes, and saturnids, but usefulness of the other groups must wait for a sounder classification. That sounder classification in turn depends, in the higher forms, chiefly on the more complete study of known characters. The matter of major cleavages, the placing of aberrant types, and especially the evo- lution of the primitive families must wait in turn for mori)hology, and partly for internal morphology. Even in the better known higher types, vastly more study of the early stages is needed. For morphology, especially internal morphology, one can find virgin terri- tory anj^vhere in the order; while in comparative physiology and the scientific study of ethology one can say nothing yet has been done. The brilliant, well-defined and well-understood pattern characters and the relatively easy breeding of the Lepidoptera make them fine objects for genetics, but so far relatively little has been accomplished. The sex mechanism, the 554 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES reverse of that in the mammals or DrosopMla, is a field for investigation. The nature of the species barrier is also a subject for study, through the various known and suspected "Rassenkreise," where one can say a population is a single species, or two, according to where in its distribution area it is studied. Goldschmidt has already studied the gypsy moth, in which the mechan- ism of sex determination is involved, but no one has touched Utetheisa (one spe- cies in the "West Indies, two from Kansas to Texas) or the buckeye (one species in Mexico, two in Florida and Cuba). Biological rather than racial speciation is an open question in Phyciodes tharos and P. hatesi, Piei'is oleracea and P. virginiensis, Halysidota tessellaris and H. harrisii, in the oleaceous-feeding or Triosteum-ieeding strains of Adita chionanthi, in the legume-feeding Thanaos haptisiae and T. afranius against the columbine-feeding T. lucilius and the sali- eaceous-feeding T. persius, and in many others. Then there is enormous opportunity for biology in the truer sense, the study of living life : natural history, life histories of individual kinds, the interaction be- tween any species and its environment, the seasons of species and strains, also behavior, and the like. This field has degenerated terribly in the last half of our century, largely, I believe, because so few people have the leisure to sit down and observe and so few live close enough to nature to be able to do so. Moreover, the study of the interaction between living creatures and their environment has become more and more sterile ever since it was christened "ecology." There are far more people who merely go through an area picking up and counting what they happen to find than there are people who know what even the commoner species are actually doing there. So much for what the next half-century can find to do. When one tries to judge what it will actually be able to accomplish, one comes to the question of means. And this seems to depend on three main items: location, leisure, and money. There are certain things that money and only money can do, and the chief of these is publication. There is always room for notes, but public sale will not finance manuals, lists, compendia, and monographs in a field of few workers like entomology. These must be financed or research will cease for lack of record of discoveries. I got a bad shock a few months ago when the announce- ment of a large amount of money for research stated inconspicuously near the bottom that this money was not available for publication. If we are to have manuals, faunal lists, and integrated surveys of biological work, the money for publication must be earmarked before the work is started and must not be di- verted. I know personally of three faunal lists that failed to appear after many hours of work, because the publication fund had vanished during the period of preparation. Also, five of the seven or eight missing volumes of Noctuidae of the Catalogue of the Lepidoptera Phalaenae, were actually prepared, but after the First World War there were no funds for printing. (I have been told that nomenclatural questions were also involved, but the part that was questioned for this reason was the only part actually published ! ) I therefore believe that the people who have control of research money, should use some of it for this type of work, and should guarantee the publication if the work is actually pre- pared in a period at all reasonable. The other necessary factor is leisure, and this is a sociological problem, for which some of the foundations bear a heavy responsibility. For they have issued HATCH: COLEOPTERA 555 masses of propaganda against the amateur, and it is only the amateur who can have the requisite leisure. There is also the question of the university, for at the moment all our intelligent young people are encouraged, and almost forced, to go to the standardized universities. These are either located in cities or have so grown and destroyed their surroundings that their students are practically cut off from contact with living nature — not merely from the Lepidoptera. We need a drive against the academic degree as a thing of high value in itself, and a restoration of the types of education that give the young adult all the things that cannot be handled in the classroom (among them living biology, as well as the fine arts, business, geography, and the like). As far as our special field is concerned, a major offender is the so-called "Graduate Record Examination," which has been getting a good deal of prestige since the Second World AVar and which, so far as I can find out, gives practically no credit score for the things a good entomologist needs to have: independent thought, aptitude, knowledge of living biology as distinguished from textbooks, and the special skills that en- able him to obtain and record his facts. And I almost forgot the museums. They are necessary recording bodies, where all the tangible and durable sources of knowledge can be preserved. They can be sources of research in only part of the field, but from every biological problem enough material should go to a museum and be saved there to enable later workers to confirm that the person who did that piece of research had what he thought he had. The museum also needs money for housing and money for care; staff for routine work and also some staff members with leisure to fol- low up research leads as they appear. This is the future for the Lepidoptera, as I see it, and equally for all fields of research in natural history. I leave it to the reader to decide how much is warn- ing and how much prophecy. COLEOPTERA Melville H, Hatch University of Washington, Seattle These remarks on a century of progress in the study of beetles may be prefaced with the caution that neither the space at the author's disposal nor his knowledge permit more than the merest synopsis of the events involved. The men and books and institutions mentioned are examples only of complex move- ments, and important names frequently may have been left unmentioned. In seeking, then, to survey the coleopterologj'- of the last hundred years, we start with men studying beetles. Beetles occur wherever men do, but different beetles occur in different regions. Tlie 3,711 species known from Great Britain (1945), the 8,473 species known from France (1935-1939), the 9,979 species and 4,409 varieties known from Italy (1929), and the 300,000 species known from the world are indices to the complexity of the problem. Two approaches to coleopterology have developed. Tlie study of local faunas has the advantage of being based on explorations that have been in month-by- 556 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES month and year-by-year contact with the insects concerned. While decades of work may be required to bring the knowledge of a local fauna to reasonable completion, the problem is of limited scope and the data are close at hand. On the contrary, the study of foreign faunas has the advantage of compre- hensiveness and of greater opportunity for general conclusions. The drawback to the broader approach is the investigator's dependence on the work of inci- dental or itinerant collectors. One is at once removed from the data, and the world as a whole is so incompletely explored that conclusions tend to lose in per- manence what they gain in comprehension. Both methods have operated together in the development of beetle studies and are, of course, strictly complementary. Those tremendous areas which lack resident collectors must be explored by the best methods available. The data obtained from the study of local faunas can be fully understood only when examined in the larger setting. It is profiitable, however, to keep the two approaches in mind as we survey the history of the past hundred years of our science. When, from the vantage point of some future century, the attempt is made to understand the development of the study of beetles, it will be seen that the hundred years just past have been part of a process of explosive develop- ment. From the perspective of a fully developed coleopterology — one that is, as a whole, at the same high level of development as the study of the German Coleoptera fauna now is the "Kaferkunde" of the present with its 300,000 known species will seem as incomplete as now appear the 594 species of Linnaeus' Sys- tema Naturae of 1758, the 22,399 species of Dejean's Catalogue of 1837, or the 77,000 species of Gemminger and von Harold's Catalogus of 1868-1876. The modern study of beetles arose in northwestern Europe, in an area roughly bounded by Great Britain, France, northern Italy, Austria, Prussia, and south- ern Scandinavia, in the mid-eighteenth century. During its first hundred years it exhibited most of the tendencies which its second century has served to con- firm and expand : the binomial nomenclature, the specific description, the de- scriptive monograph, the descriptive faunal catalogue and the faunal list, the world list, and the increasing facilities of entomological societies, journals, and musem collections. Even dichotomous analytical keys, which were first used for an entire beetle fauna in Redtenbacher's Fauna Austriaca (1849) were a product of this initial century. And in one respect this first century produced something that our second century has been unable to match, a descriptive catalogue of all previously described species, Fabricius' Sy sterna Eleutheratorum (1801), con- taining 5,172 species. Shortly thereafter the number of known species became so great that no one has since brought them together in a single descriptive work. As this first century advanced, the knowledge of beetles began to exhibit signs of maturity in portions of the area of its origin. This is shown particularly in Stephens' Illustrations of British Entomology; Mandihulata, Vols. I-V ( 1828- 1832), in which 3,650 species of British beetles were distinguished, a number that was within 100 of the 1945 figure of 3,711. There has been, of course, much reshuffling of the names in this list in the intervening century. Stephens himself reduced the count to 3,470 in his 1839 Manual, and Crotch in 1866 could list only 3,081. But the point is that this was a working out of detail. To a first approximation, the British beetle fauna had been surveyed within seventy-five years of the tenth edition of the Systema Naturae. HATCH: COLEOPTERA 557 At the same time that coleopterology was reaching maturity in its homeland, it was spreading east and west. Mannerheim (b. 1804, d. 1854) and Carl R. Sahlberg (b. 1779, d. 1860) represented the extension of beetle studies into Finland. In northeastern United States, Thomas Say (b. 1787, d. 1834) had de- scribed some 1,150 new species after 1818 and T. W. Harris (b. 1795, d. 1856) in 1833 had published a list of 994 species from Massachusetts. The greatest coleopterist of this first century of the science was Count Au- guste Dejean (b. 1780, d. 1845), peer and councilor of France. Dejean assembled the world's largest collection; he published extensively on the Carabidae of Eu- rope and the world; and he issued a Catalogue of his collections, which in its last edition of 1837 enumerated 22,399 species and was as near to a world list as the period provided. The Second Century of European Coleopterology The opening of the second century of coleopterology found the French in the ascendancy and about to produce two of the sort of sjaithetic works which are perennially necessary in an expanding empirical science, if it is to be kept from falling into chaos. The Genera des Coleopteres of Th. Lacordaire (b. 1810, d. 1870) in twelve volumes (1854—1876) — the last three volumes by F. Chapuis (b. 1824, d. 1879) — provided a description of the genera of the world. The Genera des Coleopteres d'Europe by Camile Jacquelin du Val (b. 1828, d. 1862) and L. Fairmaire (b. 1820, d. 1906) in four large volumes (1854-1868), with 292 colored plates, gave keys to and descriptions of the European genera and a synonymical catalogue of the species. At this same time W. F. Erichson (b. 1809, d. 1849), H. Schaum (b. 1819, d. 1865), G. Kraatz (b. 1831, d. 1909), and H. von Kiesenwetter (b. 1820, d. 1880) were working on the Coleoptera section of a NaturgeschicJite der Insecten Deutschlands in many volumes. Volumes I to IV appeared from 1845 to 1867, covering Adephaga, Staphylinidae, Laraellicornia, and extensive portions of the Serricornia and Clavicornia.^ Likewise incomplete and similarly ambitious was E. Mulsant (b. 1797, d. 1880) and CI. Key's (b. 1817, d. 1895) Histoire Naturelle des Coleopteres de France (1839-1884), still only fragmentary after the publi- cation of thirty-seven volumes. Not all the many-volumned faunistic surveys remained incomplete. C. G. Thomson's (b. 1824, d. 1899) Skandinaviens Coleoptera was finished in ten vol- umes, (1859-1868), and, somewhat later, AV. W. Fowler's (b. 1849, d. 1923) Coleoptera of the British Islands appeared in five volumes (1887-1891), with 180 plates, illustrating about 2,230 species. Thomson, in particular, was a very able coleopterist. He split genera rather more finely than was acceptable in his day, but ever since his names have been coming slowly into general use. The years 1862 and 1863 saw the publication of Ilagen's Bihliotheca Ento- mologica. Die Litteratur iiher das game Gehiet der Entomologie his zum Jahre 1862. A continuation of Hagen's Bihliotheca to cover the second century of Cole- optera studies is a desideratum that is only very partially met by the "Biblio- 1. The work was never completed, but later there appeared, in 1882, Vol. 11(2), by E. Reitter (b. 1845, d. 1920) on Silphidae and allies; Vol. V (1877-1920), by G. von Seidlitz (b. 1840, d. 1917) on Anobiidae and extensive portions of the Heteromera; and Vol. VI (1881-1893), by J. Weise (b. 1844, d. 1925) on Chrysomelidae. 558 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES graphia Coleotterologica Italiana" in P. Liiigioni's Coleoterri d'ltalia (1929), the "Bibliography" in Leng's Catalogue of the Coleoptera of America North of Mexico (1920), with its five supplements (1927-1948); and Musgrave's Bibliog- raphy of Australian Entomology (1932). In 1868 to 1876 there appeared in twelve volumes the Catalogus Coleopter- orum Hucusque Descriptorum of M. Gemminger (b. 1820, d. 1887) and E. von Harold (b. 1830, d. 1886), enumerating about 77,000 species for the world. In 1864 the Abbe S. A. de Marseul (b. 1812, d. 1890) established UAheille, the first journal devoted exclusively to the science of the Coleoptera. Twenty-six annual volumes of this periodical appeared to 1889. They contained mono- graphic studies, occasional French translations of German papers, biobiblio- graphical sketches of entomologists, and French translations of isolated descrip- tions of Old World beetles. After Marseul 's death, publication became irregular and finally terminated with Vol. XXXVI (1938). Marseul's idea of a journal of coleopterology was imitated. Von Harold issued the Coleopterologische Hefte, Vols. I-XVI (1867-1879). M. Cheron issued fifteen numbers of Le Coleopteriste (1890-1891); and Karl and Josef Daniel published ten numbers of their Miin- chener Koleopterologische Zeitschrift, Vols. I-III (1902-1908). The two most important extant journals of beetle studies are the Ento- mologische Blatter, Vols. 1-48 (1904-1952), founded by H. Bickhardt, and the Coleopterologische (later Koleopterologische) Rundschau, Vols. 1-31 (1912- 1948), founded by Adolf Hoffmann. Other more ephemeral serials were Pierre Lesne's Coleoptera, Vols. 1-3 (1925-1929); and Hans Wagner's Coleopterolo- gisches Centralhlatt, Vols. 1-6 (1926-1933). Adolf Horion's Koleopterologische Zeitschrift, Vol. 1 (1949) and G. Frey and Hans Kulzer's Entomologische Ar- heiten, Vols. 1-3 (1950-1952) have started publication since the war. All these journals published important contributions and monographs in their day, but the bulk of coleopterological studies appeared in journals of general entomology or of even broader scope. The second century of coleopterology had opened with coleopterists somewhat restive under the tarsomeral classification of Geoffroy (b. 1727, d. 1810). While this system indicated with some success such groups as the Heteromera or the Phytophaga-Rhynchophora complex or even the Coccinellidae, it utterly failed in the Staphjdinidae, and Erichson was seeking for ''natural" families. Dar- win's Origin of Species, 1859, opened new vistas, but it was from the penetrat- ing labors of Georg Seidlitz (b. 1840, d. 1917) of the University of Dorpat in Estonia that the modern classification really dated. Seidlitz' Fauna Baltica. Die Kdfer (Coleoptera) der Ostseeprovinzen Russlands (1872-1874) not only provided a superior faunal work for a new area, but (pp. xxviii-xxx) gave an analysis of the order into ten major subdivisions which subsequent students have done little more than rearrange and rename. Outstanding among Seidlitz' suc- cessors was Ludwig Ganglbauer (b. 1856, d. 1912), Keeper at the Imperial Nat- ural History Museum in Vienna. Ganglbauer, author of a superior but never completed descriptive catalogue of Die Kdfer von Mittelexiropa, Vols. I-IV (1892-1904), proposed in 1903- the suborders Adephaga and Polyphaga, which have since dominated most thinking along this line. 2. Systematisch-koleopterologischen Studien, Munch. Kol. Zeit. 1:271-319, 1903. HATCH: COLEOPTERA 559 There is no opportunity here to review the numerous modifications that have been suggested in the Seidlitz-Ganglbauer classification. Abdomen, wing vena- tion, male genitalia, female genitalia, head structure, larval structure have sev- erally been explored for clues regarding the natural classification of the Coleop- tera. The impression that one draws from such work is that the different special studies pretty much cancel each other out, and that the final word is still to be said.^ One of the most influential coleopterists at the middle of the second century of European coleopterology was Edmund Reitter (b. 1845, d. 1920). Beginning in 1878 Reitter wrote or edited, usually as a reprint series, the Bestimmungs- Tabellen der Europaischen Coleopteren. Some 123 numbers of this series, cover- ing most of the families, had appeared by 1942. Eventually Reitter published one of our finest beetle faunas, the Fauna Germanica, Vols. I-V (1908-1916), with 168 colored plates illustrating about 2,775 species. With its 1935 Nachtrag by Adolf Horion, it is still the standard reference work on central European beetles. Another comprehensive faunal work for an important area is Porta's Fauna Coleopterorum Italica, 5 vols. (1923-1932), and its Supplementum I, (1934), and II (1949). The classificatioin of beetle larvae is a difficult problem. Many species lead secretive lives, and their identity must usually be established in the first instance by rearing. As yet only the commoner species are known, even in Europe and America, but these are numerous enough so that diagnostic characters and keys have been worked out for the commoner genera and some of the species in most of the larger families, with the rather noteworthy exception of the Staphylinidae. Studies of larval beetles, in the period under review, started with Chapuis and Candeze's Catalogue des larves des CoUopteres (Mem. Soc. Sci. Liege, VIII : 341-653, 1855). In 1880, M. Rupertsberger (Biol. Kaf. Eur., 295 pp.) listed 1,300 European species, the larvae of which were known, a figure that he increased to 1,700 in 1894 (Biol. Lit. Kaf. Eur. von. 1880 an, 310 pp.). In 1891 W. Beutenmuller (Journ. New York Micr. Soc, VII:l-52) cited 372 North American species, the larvae of which had been noted; and in 1935 J. S. Wade {A Coiitrihution to a Bihliograpky of the Described Immature Stages of North American Coleoptera, 114 pp.) listed 1,063 species. Among the more noteworthy works on larvae were: J. C. Schi^dte's (b. 1815, d. 1884) De Metamorphosi Eleu- theratorum (1861-1888), with its 88 beautiful copper plates; M. E. Ferris' (b. 1808, d. 1872) Larves de CoUopteres (1877); A. G. Boving (b. 1869) and F. C. Craighead's (b. 1890) Illustrated Synopsis of the Principal Larval Forms of the Order Coleoptera (1931); and the section on Coleoptera in A. Peterson's (b. 1888) Larvae of Insects. Part II (1951). Recent keys to the families of larvae are found in Peterson's book and in F. I. Van Emden's "Larvae of British Beetles. III. Key to the Families" (Entom. Mo. Mag. LXXVIII :206-226, 253- 272, 1942). Meanwhile European coleopterists have been engaged in a continual assault on the foreign faunas. The museums in Paris, London, Berlin, Vienna, and else- 3. Noteworthy recent classifications are by Jeannel and Paulian, Rev. Fr. Ent. XI, 65-110, 1944; liliewise expounded by these authors in Grasse's Traite de Zool. IX : 892- 1069, 1949; and that by R. A. Crowson being published currently in the Entomologists' Monthly Magazine. 560 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES where continued to expand their collections. One major instrument of advance was the world monograph. Marseul's Histerides (1853-1861), Candeze's Ela- terides (1857-1900), Sharp's Dytiscidae (1882), Regimbart's Gijrinidae (1882- 1907), A. Schmidt's Aphodimae (1922), Jeannel's Trechinae (1926-1928), and Breuning's Carabus (1932-1937) are scattered examples of such studies. A second approach was through the study of an entire exotic fauna. Wol- laston's six volumes on the beetles of the Atlantic islands (1854-1867), and Sharp, Perkins, Blackburn, and others' six hundred pages on Coleoptera (1900- 1910), in the Fauna Haivaiiensis illustrate this sort of work. The most elaborate study of a foreign fauna is Godman and Salvin's Biologia Centrali- Americana. The Coleoptera section of this lavish work appeared between 1880 and 1910 in 18 quarto volumes, 8,703 pages, listing 18,029 species (11,675 of them new), with 350 plates (297 colored), illustrating 8,596 species. No finer monument than this exists to the British at the apogee of their imperial and industrial power. The 17 volumes on beetles (1906-1939) in the Fauna of British India, represent a partial study of another enormous beetle fauna. A third approach to the problem of the world fauna is through the study of a restricted group for a restricted area. Among the more impressive recent examples may be cited A. Hustache's Curculionides de Madagascar (1924, 582 pp.), R. Jeannel's Coleopteres carabiques de la region malgache (Madagascar 1946-1948, 1146 pp.), and P. Basilewsky's Harpalinae de'Afrique et de Mada- gascar (1950-1951, 616 pp.). P. Wytsman's Genera Insectorum (1902-1938) was an attempt — never com- pleted— to describe the genera of the world and list the species. Most of the 75 fascicules devoted to beetles were small, but the following were among the more sizable: Buprestidae by C. Kerremans (1902-1903); Elateridae by 0. Schwarz (1906-1907), Pselaphidae by A. Raffray (1908), Cicindelinae by W. Horn (1908-1915), Histeridae by H. Bickhardt (1916-1917), Aleocharinae by A. Fenyes (1918-1921), Carabinae by M. G. V. de Lapouge (1929-1932), Lagriidae by F. Borchmann (1936). The Junk-Schenkling Coleoi:>terorum Catalogue (1910- 1940, 31 vols., nearly 25,000 pp.) provided a bibliographical catalogue of about 215,000 species. Supplementa, under the editoriship of W. D. Hincks, began ap- pearing in 1950. "Winkler's Catalogus Celeopterorum Regionis Palaearcticae (1924-1932) involved the expansion of the conventional European catalogue to cover a wider area. What may be said of the new points of view that have appeared in this century in European coleopterology in addition to the continued spectacular development indicated in the foregoing paragraphs ? First, there is the greater detail of the more recent work. Darwin showed that only individuals exist, and taxonomists followed by insisting on basing their studies on ever-increasing series of specimens. Moreover, with the growth of distributional and ecological studies, there is an increasing insistance on the attachment to the individual specimens of precise data on locality, date, habitat, and collector. Second, there was the widespread tendency on the Continent to investigate infraspecific variation, with the result that large numbers of geographical vari- eties or subspecies and nongeographical varieties or aberrations have been de- scribed and named. The description and naming of very numerous non- HATCH: COLEOPTERA 561 geographical variations has occurred especially in species with variable color patterns belonging to such genera as Cicindela and Nicrophorus and to fami- lies like Coccinellidae, Ceranibycidae, and Chrysomelidae. The result was that Luigioni's catalogue of I Coleotteri d'ltalia (1929) listed 9,979 species, 1,371 subspecies, and about 3,100 aberrations in that fauna. Third, there was the discovery of the great utility of the male genitalia in the separation of species. In many genera species that can be separated only with great difficulty or not at all on the basis of external structure are readily dis- tinguished by aedeagal characters. The result has been a growing tendency in the last thirty or forty years to employ genital characters in distinguishing spe- cies, and some authors go to the extreme of regarding their figures of the geni- talia as sufficient exposition of the differences involved without supplementary verbalization. The present interest in beetles in Europe seems unabated at both the profes- sional and amateur levels. Virtually every country from the British Isles and Prance to Eumania and Russia has the requisite faunal works to aid and encourage such studies. Paulian (Col. Bull. 11:42, 1948) tells of an amateur group, the "Coleopteristes de la Seine," with more than two hundred members in Paris alone. France, Sweden, and the USSR have elaborate works on their faunas in process of current publication. In Germany there is the incredible detail of 0. Rapp's Kafer TMringens (1934-1935, 3 vols., 2,000 pages). This work starts with a list of 389 men who have contributed to the coleopterology of Thuringia. This is followed by bibliographic, distributional, and ecological data on 4,381 species, and the book concludes with an exhaustive reanalysis of the list in terms of habits and habitats. Adolf Horion, moreover, is currently issuing a new Yerzeichnis der Kafer Mitteleuropas (Abt. 1, 1951) and an ex- tremely detailed critical Faunistik der Mitteleuropaschen Kafer (Bd. I, 1941; Bd. II, 1949). The one somber note is struck by the Communist government of the USSR. Since the 'twenties the Russians have been unwilling to allow non-Communist foreigners to collect insects in their domain, and since World War II, this same prohibition has been extended to large areas in both Europe and Asia. In re- taliation, moreover, extensive areas of the West are closed to the Communists. The result is that it is impossible at present to assemble by direct field work a world collection. It is to be hoped that such conditions will not long endure. In another respect, likewise, the Russian Communists are exerting an un- fortunate influence on the study of beetles. Beginning in 1936 the Academy of Sciences of the USSR began to issue a Coleoptera section of the Faune de rURSS on a scale that promised to make it one of the great Coleoptera faunas, subjecting the beetles of the vast Russian empire to precise analysis. The first volumes were mainly in Russian, but contained extensive appendices giving German translations of the keys and of the descriptions of the new species, thus making the analysis available to an international audience. Parts issued in 1950 change this arrangement. The French title page and all sections in Ger- man, including descriptions of new species, have been eliminated, thus violating the specific recommendation of the International Code of Zoological Nomencla- ture. This vastly limits the international utility of the books and suggests that new names so proposed may be regarded as nomena nuda. 562 ^ century of progress in the natural sciences American Coleopterology in the Last Hundred Years American coleopterology first developed in the area between Washington and Boston. Say's contacts were with Dejean in Paris, and, later, LeConte and Horn were in touch with colleagues in both France and Germany. The history of the study of beetles in America was conditioned primarily by the vast hinterland which lay just beyond the Appalachian Mountains and which by 1850 extended without political or linguistic barrier all the way to the Pacific Ocean. If the situation had been different — if the Americans had been firmly hemmed in to their north Atlantic homeland or if they had been broken into several lin- guistic groups — ^American coleopterology might well have developed in accord- ance with the European pattern of increasingly detailed studies of restricted regions. Harris' 1833 list of the beetles of Massachusetts might well have de- veloped into a Massachusetts or a New England fauna. But the spell of a continent proved too strong. On the one hand, it gave a practical turn to the American mind which allowed but slight attention to such an esoteric pursuit as the study of beetles. On the other hand, it meant that such coleopterists as did appear were completely absorbed in analyzing the fauna of an entire continent. They had no energies either for the detailed local studies so conspicuous on the European scene or for the world studies which likewise, from the beginning, attracted the attention of the Europeans. The literature which did emerge took the form, almost exclusively, of technical monographs, conti- nental in scope, with the result that not many persons were attracted to the study and that American coleopterology has remained the pursuit of a few pro- fessional entomologists. The father of American coleopterology was John Lawrence LeConte, ]\I.D. (b. 1825, d. 1883), of New York and Philadelphia. A man of independent means, LeConte between 1844 and 1884 described 4,816 species of beetles in nearly all families, of which 864 were considered synonyms in 1881. Moreover, as a rule, LeConte's species were not announced in isolated publication but in mono- graphs which treated the whole continent. LeConte accompanied Louis Agassiz to Lake Superior in 1849. In 1850 and 1851 he was collecting in California and the Southwest, and in 1869 to 1872 he visited Europe, studying Kirby and Walker types in the British Museum and visiting Continental coleopterists. In 1853 LeConte joined with F. E. Melsheimer (b. 1782, d. 1873) and S. S. Haldeman (b. 1812, d. 1880) of Pennsylvania in producing a Catalogue of the Described Coleoptera of the United States, listing 4,750 species. In 1859 he edited a collected edition of The Complete Writings of Thomas Say on the Ento- mology of North America, with accompanying commentary. Since Say's col- lections had not been preserved, it was necessary to come to some understand- ing of his species as a basis for further studies. In 1861-1862 LeConte published Part I of a Classification of the Coleoptera of North America, giving the generic classification of the families except Coccinellidae, Phytophaga, and Rhyncho- phora. Part II, on Cerambycidae, appeared in 1873, but the completed work, by then revised, did not appear until a few months before LeConte's death in 1883, and then in collaboration with George H. Horn. LeConte's collection went to Agassiz' Museum of Comparative Zoology at Harvard. Of the 9,100 or 9,200 species of North American beetles known at the HATCH: COLEOPTERA 563 time of his death, LeConte had described over two-fifths. In his day he was particularly noted for his distributional studies and his suggestion that the Rhynchophora constitute one of the two primary subdivisions of the Coleoptera. In our greater perspective, it is realized that his real claim to fame is the broad descriptive basis that he laid for the study in North America of the entire order. Among the younger contemporaries and successors of LeConte who produced important monographic studies were George H. Horn (b. 1840, d. 1897), who described 1,582 new species, Frederick Blanchard (1). 1843, d. 1912), William G. Dietz (b. 1848, d. 1932), John B. Smith (b. 1858, d. 1912), Charles W. Leng (b. 1859, d. 1941), Roland Hayward (d. 1906) H. C. Fall (b. 1862, d. 1939), who described 1,453 new species, Charles F. A. Schaeffer (b. 1860, d. 1934), and E. C. Van Dyke (b. 1869, d. 1952). IVIeanwhile the indigenous study of beetles was spreading. By 1869 Johnson Pettit (d. 1898) was publishing on beetles in Ontario and Abbe Leon Provancher (b. 1820, d. 1892) in Quebec. H. G. Hubbard (b. 1850, d. 1899) and E. A. Schwarz (b. 1844, d. 1928) were at work in Detroit in the middle 'seventies, and by the late 'seventies Charles Dury (b. 1847, d. 1931) at Cincinnati and F. H. Snow (b. 1840, d. 1908) at Lawrence, Kansas, had taken up their investi- gations. The 'eighties saw John Hamilton (b. 1827, d. 1895) busy at Pittsburgh, G. W. Taylor (b. 1851, d. 1912) at Victoria, B. C, and H. F. Wickham (b. 1866, d. 1933) at Iowa City; and by the 'nineties H. C. Fall and Frank E. Blaisdell (b. 1862, d. 1947) were at work in California. Turning to synthetic works, catalogues or supplements to catalogues of Nearc- tic species have been produced in 1863-1866, 1873, 1880, 1885, 1887, 1889, 1895, 1920, 1927, 1933, 1939, and 1948. Provancher in 1877 published a Petite Faune Entomologique du Canada, Vol. I. "Les Coleopteres," describing about 950 species from Quebec and Ontario. Hamilton's Catalogue of the Coleoptera Common to North America, Northern Asia, and Europe (Trans. Amer. Entom. Soc, XVI :88- 162, 1889; ed. 2, ibid., XXI: 345^16, 1894) and Catalogue of the Coleoptera of Alaska (ibid., XXI : 1-38, 1894) were important synthesizing works. Wickham in his "Coleoptera of Canada" (1894-1899), published in parts in the Canadian Entomologist, gave keys to the species of a number of families for Ontario and Quebec. In 1910 appeared The Coleoptera of Indiana by W. S. Blatchley (b. 1859, d. 1940), which, in conjunction with Blatchley and Leng's Rhynchophora of North Eastern America (1916), provided a descriptive catalogue of 2,954 species of beetles from Indiana. Written in the tradition of LeConte and Horn, with its division into Genuina and Rhynchophora, this is the only complete descriptive beetle fauna, except Provancher's provisional work, produced so far in North America. Leng's catalogue of the Coleoptera of America North of Mexico (1921), broke with the LeContian tradition and integrated American studies with those that had been going on in Europe. M. H. Hatch's (b. 1898) Indices to keys and local lists (1927-1928) (Journ. New York Entom. Soc, XXXV :279-306, 1927; XXXVI :335-354, 1928; XXXVII :135-143, 1929; XLIX:21^2, 1941) organ- ized aspects of the literature, and J. C. Bradley's (b. 1883) Genera of Beetles of America North of Mexico (1930) provided a much-needed revision of LeConte and Horn's Classification. 564 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Between 1884 and 1924 Thomas L. Casey (b. 1857, d. 1925) described some 9,400 species, mostly from the Nearctic region. Large numbers of these were based on evanescent differences and are invalid by conventional criteria. W. Horn (1915) rejected 86 out of 99 of Casey's names in Cicindelidae; Leng (1920) re- jected 144 out of 150 names in Buprestidae and 30 out of 34 names in Prionini; Banninger (1950) could recognize only one out of 24 names in Pasimachus; and Karl Holdhaus (Schroder's Handh. d. Entom. 11:899, 1927) complained that Casey had so multiplied species in numerous families as largely to conceal the true status of the Nearctic fauna. Casey left his collection to the National Mu- seum, and, whatever one thinks of Casey's work, there can be nothing but praise for the generous, intelligent cooperation shown by Mrs. Casey and the officials of the Museum in preserving his material for future students. The almost exclusive preoccupation of Americans with their own fauna has already been noted. G. H. Horn covered Throscidae and Eucnemidae in 1890 for the Biologia Centrali- Americana; A. Fenyes (b. 1863, d. 1937), of Pasadena, covered Aleocharinae for Genera Insectorum (1918-1921); and M. H. Hatch, of Seattle, did the Silphidae (1928) and Leiodidae and Clambidae (1929) for the Coleopteronim Catalogus. But such contributions only served to emphasize the general absence of Americans from the international scene. American interest in the Neotropical fauna was signalized in 1914 when Leng, in collaboration with A. J. Mutchler (b. 1869), of the American Museum, pub- lished a List of the Coleoptera of the West Indies (Bull. Amer. Mus. Nat. Hist., 33:391^93). W. S. Fisher (b. 1878), at the United States National Museum, revised the West Indian Buprestidae in 1925 (Proc. U. S. Nat. Mus., 65(9) :1- 207) and he and other coleopterists at that institution displayed a persistent in- terest in the Neotropical fauna, which culminated in R. E. Blackwelder's (b. 1909) Monograph of the West Indian Staphylinidae (Btdl. TJ. S. Nat. Mus., No. 182, 1943) and Checklist of Neotropical Coleoptera, 1944-1947 {Bull. TJ. 8. Nat. Mus., No. 185). The opening up of automobile communication with Mexico in the 'thirties and other factors stimulated contacts with the south, which have re- sulted in contributions to the knowledge of the Neotropical fauna by Orlando Park (b. 1901), of Northwestern University, in Pselaphidae, E. G. Linsley (b. 1910), of the University of California, in Cerambycidae, M. A. Cazier (b. 1911), of the American Museum, and numerous others. World War II similarly served to broaden American horizons so that, for instance, P. J. Darlington (b. 1904), of Harvard, is devoting himself to circumtropical Carabidae. The limited status of coleopterology in the United States is revealed by the fact that a year after its founding in 1949 a Coleopterists' Society had only 186 members, and died in 1952. The largest group of coleopterists in the country is at the National Museum in Washington, where E. A. Chap in (b. 1894) is curator and where the economic importance of the order assures the continua- tion of a staff interested in beetles. Similar economic motives also assure the permanence of such studies at Ottawa, where W. J. Brown (b. 1902) has been in charge of Coleoptera since 1927. Sizable groups of coleopterists are likewise centered at the Chicago Museum, where R. L. Wenzel (b. 1915) is curator, and in the San Francisco Bay area, where E. S. Ross (b. 1915) is curator at the Cali- fornia Academy of Sciences and E. G. Linsley chairman of the Department of Entomology at the University of California. This last institution, in particular, HATCH: COLEOPTERA 565 since well before E. C. Van Dyke's retirement in 1939, has trained a notable series of able eoleopterists. In 1947 E. H. Arnett (b. 1919), then at Cornell University, founded The Coleopterists' Bulletin, the first American journal of coleopterology. COLEOPTEROLOGY IN AUSTRALIA, NeW ZEALAND, AND ELSEWHERE Indigenous coleopterology in Australia dates from the founding of an Ento mological Society of New South Wales in Sydney in 1862 by Sir William Mac- leay (b. 1820, d. 1891), the Reverend R. L. King (b. 1823, d. 1897), and others. George Masters (b. 1837, d. 1912) published a Catalogue of the Described Cole- optera of Australia in 1871, with a second edition in 1885-1887 and a supple- ment in 1895-1896. The Reverend Thomas Blackburn (b. 1844, d. 1912) de- scribed 3,069 species of Australian beetles between 1888 and 1912. Other note- worthy students of Australian Coleoptera have been T. G. Sloane (b. 1858, d. 1932), a specialist on Carabidae; A. M. Lea (b. 1868, d. 1932), of the South Aus- tralian Museum at Adelaide, and H. J. Carter (b. 1858, d. 1940), who deposited his collection in the National Museum at Melbourne. A. Musgrave (b. 1895) published a useful Bibliography of Aust7'alian Entomology in 1930. In 1926 there were 16,660 species of Coleoptera known from Australia. The indigenous study of New Zealand Coleoptera is virtually the work of one man. Major Thomas Broun (b. 1838, d. 1919), of Auckland. During thirty- nine years preceding his death Broun described some 4,323 species from the archipelago, of which about 3,500 were named by him. G. V. Hudson in his in- teresting book, Neiv Zealand Beetles (1934), suggested that perhaps as many as half of Major Broun 's species may be synonyms ! The author of this paper lacks the knowledge, even if he were permitted the space, to attempt a country-by-country evaluation of Coleoptera studies. South- ern South America is the site of some work, of which Carlos Bruch's (b. 1869, d. 1943) Catdlogo Sistemdtico de los Coleopteros de la Repiiblica Argentina (1911-1914), with Suplemento I-IV (1915-1928), is an outstanding example. South Africa is likewise the home of some indigenous study of beetles, the Bel- gian-born L. A. Peringuey (d. 1924) having been one of the leading contributors. Indigenous coleopterology developed late in Japan, where most of the species have been described by Europeans. H. Kono, T. Kano, and Y. Miwa are out- standing among Japanese coleopterists, the last being the author of A System- matic Catalogue of Formosan Coleoptera (1931). In China, before the Com- munist revolution, a certain amount of coleopterological work was under way, especially at some of the Japanese- and American-sponsored institutions. Y. Ouchi's Biographical Introduction to the Study of Chinese Insects (1934), C. F. Wu's Catalogus Insectorum Sinensium, Vol. 3, "Coleoptera" (1937), and J. L. Gressitt's Longicornes de Chine (1951) are examples. In Hawaii, E. C. Zim- merman (b. 1912) has been studying the beetles of Oceania since 1934. Fossil Beetles The early students of fossil beetles, like 0. Heer (b. 1809, d. 1883) and S. H. Scudder (b. 1837, d. 1911), assigned the remains with great exactness to living ^^(, A CENTURY OF PROGRESS IN THE NATURAL SCIENCES genera. By 1908 some 352 species of Mesozoic and 2,559 species of Cenozoic beetles had been described, nearly all from Europe and North America. Beetle fossils, however, consist almost exclusively of impressions of elytra and pronota. In view of the slight extent to which characters drawn from these parts are de- cisive in the classification, A. liandlirsch (b. 1865, d. 1935) in his Fossilen In- sekten (1908) suggested that, so far as the Mesozoic remains were concerned, the record revealed no more than the presence of certain general coleopterous types, such as carabid, elaterid, buprestid, and hydrophilid. The Tertiary re- mains, however, even as far back as the Eocene, continue to be placed in such living genera as Lehia, Harpalus, Bemhidion, Platynus, Berosiis, Tropisternus, Bledius, Lathrohium, Cryptocephglus, and Sitona. So far as the elucidation of interglacial and postglacial remains of beetles is concerned, a good deal of progress is being made by a more intensive study of the characteristics of the elytra and pronotum of living species. Much further back in the geological record than this, however, the writer feels that coleopter- ists probably must be satisfied with form-genera, many of which may never be integrated satisfactorily in the classification of living types. In 1924, Tillyard (Proc. Linn. Soc, New South Wales, 49:429-435, 1924) described six species of Coleoptera from the Upper Permian of New South Wales along with a beetlelike wing cover exhibiting true venation, which he ascribed to a new order, Protoeoleoptera, close to the ancestry of the Coleoptera. A. V. Martynov (b. 1870, d. 1938) in 1933 reported beetles from the Permian of Russia, and Jeannel (1947) erected a suborder Archicoleoptera for these Permian beetles. Jeannel, furthermore, lias shown considerable enthusiasm for the Gondwanaland hypothesis and Wegener's "wandering continents" as an aid in understanding beetle distribution, but there are many who disagree with him. Prognostication of the future of coleopterology is uncertain. If it follows the pattern of a relatively mature science like ornithology, the time will come when beetles everywhere, in both adult and larval stages, will be as well known as are now the adult stages of central and northwestern Europe. The geographi- cal variability and the ecological relationships of the species, likewise, will be worked out. Many years will be required to realize such a program ! STREPSIPTERA R. M. BOHART University of California. Davis The status of knowledge in the order Strepsiptera in 1853 is indicated by the fact that only 5 of the 23 currently known genera had been described and these represented only 3 of the 5 families as we now know them. Some two dozen authors had contributed descriptions and figures dealing with a total of 11 species but had devoted most of their publications to a discussion of the phylogenetic relationships of these peculiar insects. The first strepsipteran was BOHART: STREPSIPTERA i 567 described by Peter Rossi in 1793 and twenty years later, in 1813, Kirby erected the order Strepsiptera. This did not find favor witli the majority of taxonomists, and the group was placed generally in the Coleoptera or less frequently in the Diptera, Neuroptera, or Hymenoptera. In the following fifty-five years only 3 more genera were described but our knowledge was increased to a greater extent along other lines. The only fossil strepsipteran now known, Mengea tertiaria Grote, was reported by Menge in 1866 in Baltic amber. This indicated that the order had not changed greatly since Tertiary times. In 1893 Nassonow gave the first account of internal anat- omy and his work is still the best available on the subject. Another step forward was a paper by Perkins in 1905, which described the life histories of several parasites of leafhoppers and suggested their possible importance in biological control. The stage was set for an expansion of the Strepsiptera along systematic lines, and from 1909 to 1918 W. D. Pierce dominated this field. Holding stubbornly to a theory of host-parasite specificity, Pierce raised the number of described species by 1918 to a total of 166, which he distributed among 5 superfamilies, 11 families, 8 subfamilies, 5 tribes, and 49 genera. Fortunately, later workers have been able to reduce this top-heavy structure to 6 families and 16 genera, to which 7 more have been added. The chief value of Pierce's papers was that they assembled for the first time all the scattered references to the order so that workers in various parts of the world could proceed on a common basis. At this time (1918) it was thought that all female Strepsiptera were endo- parasitic for life, beginning with the second larval stage. This idea was blasted by Peyerimhoff, who described the free-living female of Eoxenos in 1919. An- other major contribution was that of Salt (1927) dealing extensively with the effects of stylopization. He pointed out that in the Hymenoptera those hosts which had a fixed amount of larval food, such as the solitary Vespidae, fre- quently assumed the characters of intersexes, whereas the hosts continually fed as larvae, such as the social Vespidae, exhibited no such external differences. The work of Peyerimhoff bore fruit after fifteen years when, in 1934, Parker and Smith associated the female Eoxenos with a mengeid male and established the female of the Mengeidae for the first time. All that remained to complete the skeleton framework of the picture in this family was to find the host. This turned out to be a thysanuran, as reported by Carpentier in 1939. In the same year Ogloblin published the first evidence of females in the family Myrmeco- lacidae. His startling finds indicate that the males parasitize ants and the females mature in various types of Orthoptera. Also in 1939, Lindberg, working with Elenchus parasitic on a fulgorid, gave the first complete record of the re- lationship between a strepsipteran and its host. The first major review of the order since that of Pierce in 1918 was attempted by Bohart (1941) who revised the world genera and the species of North Amer- ica. It was here that many of the superfiuous categories of Pierce were synony- mized. Bohart followed this with the first comprehensive paper on the leafhopper parasite family Halictophagiclae in 1943. Also in 1943, Hofeneder and Fulmek published a complete cross-reference catalogue to the parasites and their hosts, and Silvestri gave a comprehensive treatment of the biologies of 6 Italian species of Mengenilla. 568 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES At present there are only a few workers actively publishing papers on the Strepsiptera. Certainly one reason for this is the difficulty of getting enough material for study. Male specimens, which offer the best characters for syste- matic study, are generally rare. Exceptions to this rule have been pointed out by MacSwain (1949), who collected 510 male Stylops by exposing 8 caged fe- males, and by Bohart (1951), who reported nearly 300 male myrmecolacids taken at light in the Philippines by E. S. Ross and by H. Hoogstraal. Although the idea of complete host-parasite specificity has been largely dis- credited, the controversy with respect to the phylogenetic position of the order still continues. Probably the majority of systematists favor the ordinal status but a strong minority would place these insects with the Coleoptera, and a dis- sident few attempt to relate them to whiteflies and scales. REFERENCES Bohart, R. M. 1941. A revision of the Strepsiptera witli special reference to tlie species of North America. Univ. Calif. Publ. Entom., 7:91-160. 1943. New species of Halictophagus with a key to the genus in North America. Ann. Entom. Soc. Amer., 36:341-359. 1951. The Myrmecolacidae of the Philippines. Wasmann. Journ. Biol., 9:83-103. Carpentier, F. 1939. Sur le parasitisme de la deuxieme forme larvaire d'Eoxenos lahoulienei Peyer. Bull, and Ann. Soc. Entom. Belg., 79:451-468. HoFENEDER, K., and L. Fulmek 1942-1943. Verzeichnis der Strepsiptera und ihrer Wirte. Arb. physiol. agnew. Entom. Berlin-Dahlem, 9:179-185, 249-283; 10:32-58, 139-169, 196-230. KiRBY, W. 1813. Strepsiptera, a new order of insects. . . . Trans. Linn. Soc. London, 11:86-123. LiNDBERG, H. 1939. Der Parasitismus der auf Chloriona-Arten lebenden Strepsiptere ElencMnus chlorio7iae n. sp. sowie die Einwirkung derselben auf ihren Wirt. Acta Zool. Fennica, 22:1-179. MacSwain, J. W. 1949. A method for collecting male Stylops. Pan-Pac. Entom., 25:89-90. Menge, a. 1866. Ueber ein Rhipidopteren und einige andere im Bernstein eingeschlossene Tiere. Schr. Naturf. Ges. Danzig, ser. 2, 1 : 3-4. Nassonow, N. V. 1893. On the morphology of Stylops melittae. Warsaw Univ. News (1893), pp. 1-30. (In Russian.) Ogloblin, a. a. 1939. The Strepsiptera parasites of ants. 8th Internat. Congr. Entom. Berlin, Ver- handl., 2:1277-1284. Parker, H. L., and H. D. Smith 1934. Further notes on Eoxenos lahoulhenei Peyerimhoff, with a description of the male. Ann. Entom. Soc. Amer., 27:468-479. BROWN: ANT TAXONOMY 569 Perkins, R. C. L. 1905. Leafhoppers and their natural enemies. Hawaiian Sugar Planters' Assn. Exper. Sta. Bull., 1:90-111. Peyerimhoff, p. 1919. Un nouveau type d'insectes Strepsipteres. Bull. Soc. Entom. France, 1919: 162-173. Pierce, W. D. 1909. A monographic revision of the twisted-winged insects comprising the order Strepsiptera Kirby. Bull. U. S. Nat. Mus., 66:1-232. 1918. The comparative morphology of the order Strepsiptera together with records and descriptions of insects. Proc. U. S. Nat. Mus., 54:391-501. Rossi, P. 1793. Observation de M. Rossi sur un nouveau genre d'insecte, voisin des Ichneumons. Bull. Soc. Philom. Paris, 1:49. Salt, G. 1927. The effects of stylopization on aculeate Hymenoptera. Journ. Exper. Zool., 48:223-331. SiLVESTRI, F. 1943. Studi sugli "Strepsiptera" III. Descrizione e biologia di 6 specie italiane di Megenilla. Boll. Lab. Zool. gen. agrar. Fac. Agrar. Portici, 32:197-283. ANT TAXONOMY W. L. Brown, Jr. Museum of Comparative Zoology, Harvard University When the subject, title and dates for this survey were suggested to me, I could not but be struck by the coincidence of the century now complete with what I see as a major, if not entirely progressive, phase in the development of ant systematics. If we begin the history of our myrmecological century at 1853, we find al- ready existing a primitive chaos of Linnaean binomials scattered among "For- mica" and a few other form genera. In the heyday of the two-line Latin diag- nosis, beginning with Linnaeus in 1758, many workers in the general field of insect taxonomy — Fabricius, Latreille, Westwood, and otliers like them — ac- cumulated a great many species of ants as mere incidents to their systematic outpourings. Only Latreille, and toward the close of the period, Nylander, gave the taxonomy of the ants more than a passing glance. In the few years closely centered on 1853, three men, Gustav Mayr, Julius Roger, and Frederick Smith, entered the scene with publications focused more or less directly and exclusively upon ants. The study of the family would un- doubtedly be farther advanced today had Smith never chosen to look at an ant, but this has been emphasized by so many authors already that I hardly need labor the subject. Creighton, in the historical introduction to his The Ants of North America covers Smith's work adequately and, in my opinion, with considerable restraint. At one point, he paraphrases Forel as stating "with his characteristic impetuosity . , . that neither Smith's species nor his types could 570 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES be depended upon," and continues, "but while Forel's annoyance is understand- able, he obviously overshot the mark. It is only because of Smith's types . . . that Smith's work was saved from oblivion." For myself, after considerable ex- perience with Smithian species and Smithian "types" in the British Museum, I can only side with Forel on this question. Smith himself, and a few of his con- temporaries and successors in the British Museum, had a genius for mistran- scription, label-switching, and outright substitution or loss of specimens that has seldom if ever been equaled in the history of entomology. While it is no longer necessary to add to the damnation of Smith's work, it is important that the authenticity of his types remain open to question. Passing to Roger, we find a man of a different stamp. His publications are relatively few in number, but the descriptions are very thorough for his day. He had a remarkable eye for genera. He struggled, as did Mayr, with the con- fused inheritance from the Linnaean period and with Smith's descriptive atroci- ties, and he reduced a goodly share of the mess to ordered synonymic lists. In all his work, Eoger showed caution and restraint in the face of the exciting bizarre novelties then appearing in Europe from the corners of the world. In Gustav Mayr, we come to a truly great myrmecographer. While main- taining other interests, he gave his best attention to the ants. Like Roger's, his descriptions were meaningful and perhaps even more to the point. Mayr early tackled the most important problem then confronting ant systematics — the gen- era and higher categories. What needed doing then is obvious to us now largely because Mayr did it. Starting with the Palearctic fauna, and then taking on the exotics, he apportioned with great insight the known and new forms into the familiar genera we know today, carefully characterizing each genus as he went. Fighting to recognize and correctly place not only the tremendous back- log of old species but also the spate from Smith's activity, he nevertheless found time to describe a great many species with a clear sense of the significant charac- ters and a conservative approach to intraspecific variation that present-day in- vestigation is ever more solidly confirming as superior to the fine nomenclatural splitting practiced by most of his successors. Mayr's names largely stand today as steady reference points in the taxonomic maze. About 1870, in the middle of Mayr's course, Emery and Forel started their prolific taxonomic careers. The parallels and divergences between their lives and work has been covered by Creighton. Both Emery and Forel began with modest and useful studies of the European fauna, and Forel completed studies of great importance in his early publications on the comparative anatomy of the gizzard, poison apparatus, and anal glands, recognizing most of the features still serving to distinguish the major subfamilies. Forel, however, soon discovered the unlimited taxonomic possibilities of the vast collections of ants rapidly accumulating in Europe with the develop- ment of the colonial empires. His work on ants then largely settled down to a routine of descriptions of exotic collections, one by one, and the numbers of species, subspecies, and varieties bearing his name rose steadily into the thous- ands. Creighton's estimate of Forel's descriptive efforts, while largely critical, is surprisingly mild, perhaps owing to the relatively small role played by Forel in the description of North American Formicidae. Even this role, as repeatedly shown in the synonymy of Creighton's book itself, was not a particularly dis- BROWN: ANT TAXONOMY 571 tinguished one. Creighton's claim that Forel described ants "with ability and distinction," and his estimate that "among the great number of new ants which [Forel] described comparatively few were synonyms" are concessions too charit- able for me to accept without protest. After his promising start, Forel's taxo- nomic career was one protracted degeneration into ever more hasty, careless, and often pointless proliferation of new names. I doubt very seriously that the year 2053 will see as many as one half of the names proposed by Forel in good taxo- nomic standing. Forel undoubtedly had a highly developed intuitive knowledge of the distinctness and affinities of many of the ants with which he dealt, and it is myrmecology's loss that he did not often pause long enough in his headlong pursuit of new forms to make clear either their distinctive characters or their real relationships. Excessive hurry, looseness, and confusion are the obvious marks upon most of Forel's publication, and the pentanomial system his charac- teristic medium of taxonomic expression. Carlo Emery approached Forel in numbers of species described, and sur- passed him in genera. In his early years, he produced a number of very useful papers, now all but forgotten, in which dozens of names from the old inquiren- dae lists were hunted down and tucked safely into the synonymic structure. His descriptions were more pointed than Forel's, and usually much more precise; many of the abundant illustrations he furnished, while often inaccurate in de- tail, provide the best evidence as to what the species of Smith and Forel are really like. Emery spent a large part of his physically handicapped career in the attempt to revise, classify, and key the species, genera, and higher categories, and in his classic contributions to Wytsman's Genera Insectorum he produced a unified system, key, and complete catalogue of the ants — the most useful work published in myrmecology to date. With Forel, he followed the weird and won- derful pentanomial system, but utilized it with much greater moderation than did Forel when describing novelties. Emery worked well and conscientiously, but the flood of unreliable contemporary description hurried him too much and threw him off the track at important junctures in his classificationary labors. Curiously, and unlike ]\Iayr, Emery seems to have expressed remarkably little criticism of the work of his contemporaries, even though the constant inter- change of types with Forel, Santschi, Wheeler, and others must have alerted him to their inconsistencies. It was calamitous that these authors should have been allowed to publish so copiously and for so long without the critical check earlier exercised by Eoger and Mayr on the woi'k of Smith. Only late in his life does Emery seem to have realized the extent of the damage done, as is ap- parent in his angry but flagging attacks on feckless dabblers like Bondroit and Donisthorpe. W. M. Wheeler entered the field in 1900, and within a few years produced the general text, Ants, still in use but badly outdated. Wheeler's taxonomic writings came thick and fast, and were similar in style and quality to those of Forel, except that they were more frequently accompanied by illustrations and keys and were often weighted according to biological information gained in the field. Wheeler's work, like that of Forel, declined seriously with advancing years. His best contributions to taxonomic myrmecology were, perhaps, his studies on ant larvae and his treatment of the Baltic Amber fauna. The years following 1910 saw many specialists joining the rush to describe 572 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES ants: Santschi, Kuzsky, Stitz, Viehmeyer, Karawajew, Bondroit, Donisthorpe, Crawley, Menozzi, Clark, and numerous others. Taken generally, their work is very disappointing, following as it does more or less faithfully the pattern of Forel in spirit and method. One looks in vain among the thousands of dubious names and useless descriptions published by these workers for a real sign of a developing critical approach, but all that meets the eye is "sp. nov.," "subsp. nov.," "var. nov.," punctuated very occasionally by an irrelevant figure or an unworkable key. The freshest works of the period are probably those of Arnold and Mann, based on material largely collected by themselves in relatively remote and myrmecologically unknown parts of the world, and produced as whole faunas with keys and figures. The reaction to this depressing period of description for description's sake and increasing taxonomic irresponsibility was dreadfully slow in gathering strength. In the 1920's and 1930's, the center of myrmecological investigation began, almost imperceptibly at first, to shift from Europe, with men like Bruch, Gallardo, and Borgmeier in South America, Arnold in South Africa, and M. R. Smith and Creighton in North America concentrating more closely upon the native ants of their own regions. In the light of their field observations and careful collecting, the pentanomial system came under a severe strain, and at the same time there arose a feeling that the art of description had fallen to a very low state. Improvements in techniques of sampling, description, and illus- tration became general, in large part at the insistence of Kennedy, but it was not until the appearance, in 1942, of Ernst Mayr's Systemaiics and the Origin of Species that the stage was set for the loosening of the debilitating grip of the pentanomial system upon ant taxonomy. This grip was first broken for myrme- cology by W. S. Creighton's Ants of North America, appearing in 1950, a book that not only applied Ernst Mayr's principles broadly to a large fauna but finall}' signaled an uncompromising shift to the critical, revision-minded, bio- logical taxonomy we hope is here to stay. After three years, it seems certain that Creighton's book is having a resounding effect on taxonomic theory and practice around the world, and it is especially gratifying to note that the younger work- ers are approaching the study with a revisionary spirit. Because Emery's and Wheeler's generic keys are based on an unsound system to begin with, and because they have been swamped by the description of the past thirty years, the outstanding need in general ant taxonomy today is a new and workable key to the genera and higher categories. This must be based on a new and sounder classification, which in turn requires dehridement through wholesale synonomy at all systematic levels and a thorough survey of compara- tive anatomy, both external and internal, in the various ant groups. Modern generic revisions, thoroughly done, deserve and are now receiving high priority. A survey of the male genitalia is badly needed. A look at recent publications and work in progress today shows a response to these needs that is encouraging on the whole, and there seems to be no reason why the current gratifying trend should not continue. Because of their huge and readily available populations and their segregation into colonial systems capable of considerable manipula- tion, ants provide a marvelous kind of material for biological study. It would be a shame if the taxonomic picture were to remain so confused as to continue seriously to hamper their usefulness. HURD: THE ACULEATE WASPS 573 THE ACULEATE WASPS Paul D. Hurd, Jr. University of California, Berkeley In any review of the work accomplished during a certain period of time what we really are doing is attempting to examine accumulated knowledge in the light of the present in order that we may from the empirical evidence project our lines of thought toward the future. It is helpful, tlierefore, to evaluate the nature of the work undertaken in the study of the aculeate wasps during the past century. Fortunately for the purposes of establishing a natural point of reference, Frederick Smith published between 1853 and 1859 a catalogue of the hymenopterous insects in the collection of the British Museum, which, in a measure, not only provided a summary of the knowledge of the known wasp fauna of the world at that time, but, more importantly, pointed up the nature of the investigations which had preceded this date. Large areas of the earth's surface were unexplored. Those areas that had received the attention of the hymenopterist were so poorly known that even a guess as to their faunistic composition and relationship could not be safely hazarded. The classifications of earlier writers (mainly those of Latreille, Le- peletier, and Dahlbom) were to a large degree inadequate and failed to afford a true reflection of the nature and extent of the world wasp fauna. To be sure, the wasp faunas of certain major political districts, such as England, France, and Germany had received more intensive study and were accordingly consid- ered comparatively well known. Several important lines of investigation suggested themselves following the appearance of the catalogue. Perhaps paramount was the realization that much material would be needed from Africa, Asia, Australasia, and the New World before a better understanding of the world wasp fauna would be forthcoming. Study of materials from the more poorly explored regions of the world sug- gested that much revision of ideas concerning relationship, distribution, and biology would be necessary. Consequent on these needs a greater effort to ac- quire faunal representatives from the large biogeographical regions of the earth was manifested in the increasing number of scientific expeditions. So remark- able were some of the discoveries in foreign lands that travelers and voyagers would return with tales of the gigantic sizes of the wasps. From the 1850 's until after the turn of the century the results of many of the exploratory expeditions were reported upon. The work of this period largely centered about the description of the material acquired and was usually in the form of large faunal works covering continental or subcontinental regions. Note- worthy contributions on this scale were made by Andre in Europe and North Africa (1882 et seq.), Ashmead in Hawaii (1901), Bingham in India (1897- 1913), Cameron in Central America and the Orient (1888 et seq.), and Cresson (1867 et seq.) in North America. Toward the end of the nineteenth century an important deviation in the type of treatment occurred. Faunal studies began to be reduced in geographic scope. The revision or monograph of various cate- gories usually of generic or familial level became more popular and provided a 574 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES better method of analyzing and making known the composition of the better- collected faunal districts of the world. As each increment of information of a particular group or region was added to the fund of knowledge, the total bio- geographic picture commenced to emerge. In 1882 Alexandro Mocsary published a comprehensive world list of the lit- erature pertaining to the order Hymenoptera. This was followed a few years later by the appearance of Dalla Torre's Catalogus Hymenopterorum (1892- 1902), a work which provided a stimulus for the more exhaustive monographic treatments which were to follow. More attention began to be directed toward accumulating more detailed information on the distribution and biologies of certain groups of wasps. Unfortunately little effort seems to have been made toward tying together all the available information on any one group. As new frontiers of the world were opened, largely through improved methods of trans- portation, so many new species were being collected that the taxonomist devoted a large share of his time to providing names. In America, Thomas Say was chiefly responsible for initiating the descriptive phase in this country. Ezra T. Cresson (1863) brought together in his catalogue the described species of North American wasps. Cresson led the way in com- mencing an exhaustive study of the wasp fauna of North America. Similar in- vestigations had preceded these — principally in England, France, and the Ger- man countries. The results of the European studies, as well as the influence of their workers, largely guided American thinking in matters of classification, phylogeny, and biology. By 1887 Cresson had presented a synopsis of the North American families and genera. At the turn of the century Ashmead re-examined the existing classifications and made an attempt to synthesize the existing knowl- edge relating to the phylogeny of the Hymenoptera. Other workers, such as Viereck in America, Andre in France, Cameron in England, and Bischoff in Germany, began to shape the broad outlines of the next twenty-five years of re- search on the wasps. In general, the lower categories, particularly on the generic level, were accorded a more thorough and virtually monographic treatment. This approach was, to be sure, closely correlated with advances in the related sciences and the improved technological equipment at their disposal. Perhaps the most significant contribution to the knowledge of the aculeate wasps made during the present century has been the application of the prin- ciples stemming from the theory of evolution. While it is yet too early to deter- mine the total effect this will have on the analyses and evaluations of problems dealing with biogeography and phylogeny, it is apparent that it will be profound. The present trend of study has assumed the form of synthesis of the various branches of knowledge so that the emerging interpretation of the aculeate fauna is directed toward reflecting the equivalency expressed in nature. This method seems best to achieve the ideal representation of the facts concerning the origins, phylogenies, ecologies, and the role of the wasps in nature. In order to accomplish this interpretative representation it might be w^ell first to re-examine more closely the present outlook on the basis of the probable world aculeate fauna. The recent catalogue of Nearctic Hymenoptera lists ap- proximately 3,500 species and infraspecifics from an area representing nearly one-sixth of the earth's surface. Allowing for compensating changes in status, synonymy, and description of new species, as well as taking into consideration MICHENER: THE APOIDEA 575 the relative ecological iinequalness of faunas there are probably no more than 20,000 species of aculeate wasps inhabiting the surface of the earth. A figure of 15,000 species is more likely nearer the actual number, especially when infra- specific categories are taken into consideration. At first glance this figure seems small and suggests the possibility that at least certain main aspects of the study of the total world aculeate wasp fauna may soon be realized. This is particu- larly encouraging, for if we multiply 15,000 fourfold in an attempt to gain an appreciation of the principal developmental stages requiring morphological de- scription alone, the chore ahead of us seems proportionately greater. Com- pounded to this arithmetical evaluation of extrapolated progress are those in- tangible aspects of the study which involve the fields of evolution, physiology, economics, and so forth, which, if reduced to a numerical power of 15,000 sug- gest an almost hopelessly astronomical figure — indeed one unobtainable in the life expectancy of the earth if presently employed methods of research and recording remain essentially the same. The studies of the past century have provided us with a fund of knowledge — largely unsynthesized and scarcely subjected to interpretation — a basis, as it were, for theoristic advances in thinking and methodology so as to guide us in our ideal representation of the world wasp fauna. THE APOIDEA Charles D. Michener University of Kansas Taxonomy The period 1853 to 1953 is particularly appropriate for a review of our knowledge of bees because in the year 1853 Part I of Frederick Smith's Cata- logue of Hymenopterous Insects in the Collection of the British Museum ap- peared. This and the next part of the same work, published in 1854, dealt with the bees. In these publications a vast number of genera and species from all parts of the world were described. The first step in making known information on any group of organisms has always been the naming of the species involved. Numerous previous authors had started this process, so that most of the bee species of Europe were known by 1853 (see, for example, Kirby's Monographia Apum Angliae, 1802) and numerous species from elsewhere had also been de- scribed. The most comprehensive descriptive work prior to 1853 appeared in 1836 and 184:1— Histoire naturelle des insectes, Hymenopteres, by Lepeletier de Saint^Fargeau. For their time both Lepeletier and Smith did excellent work, which has served subsequent bee students as well as can be expected. From Smith's time to the present there has been a continuous series of au- thors describing species of bees from various parts of the world. In this country E. T. Cresson, of Philadelphia, described a great many bees, most of them in the years 1878 and 1879. Curiously, although Cresson lived and worked in Penn- 576 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES sylvania, most of his studies were based upon specimens brought back by numer- ous collectors in the West, for the scientific exploration of western America was in full swing in Cresson's time. Many of the bee species which could be collected in Philadelphia itself went undescribed during Cresson's activities and it re- mained for Charles Robertson to discover and name them in Illinois, mostly during the last decade of the last century and the first decade of the present one. Probably because of the activities of Cresson and Robertson in this country and of I'Abbe Provancher in Quebec (many of whose species were incorrectly placed generically and still remain to be elucidated), European authors avoided work on North American forms after Smith's time. They studied material from all other parts of the world, and H. Friese in particular described thousands of species from all faunal regions except the Nearctic. His work extended over a very long period, at least from 1891 to 1935. Friese's counterpart in America was T. D. A. Cockerell, who first published on bees in New Mexico in 1894 and whose last work, on bees from Honduras, appeared in 1949. During this long period Cockerel described bees from all parts of the globe, and he himself col- lected them in many countries. In addition to the publications of these workers who have studied bees from all parts of the world, notable contributions in collecting and naming bee species have been made by a number of students whose interests or opportunities have been more localized, for example Tarlton Payment in Australia, E. L. Holm- berg, Padre J. Moure, and C. Schrottky in South America, and P. H. Timber- lake in California. Others have specialized on certain groups of bees, and have often contributed more of lasting value than those whose work has been of a faunal nature. Examples are II. J. Franklin (Bombini), T. B. Mitchell {Mega- chile), P. Bliithgen (Halictinae), and H. F. Schwarz (Anthidiini and Meliponini) . The result of all this activity has been a very large number of described species of bees. In the recent catalogue of Hymenoptera of America North of Mexico by C. F. W. Meusebeck et ah, 3,285 species and subspecies of bees are listed. Some of these will prove to be synonyms, but at least as many new ones will probably be described. Assuming that there may be 4,000 species in the entire North American continent and that the other major continents (South America, Eurasia, Africa) average 4,000 additional species each, while in Australia and insular regions another 3,000 species exist, we reach a total of 19,000 species. This is remarkably close to an estimate of 20,000 made many years ago by Friese. Phylogeny As large numbers of bee species were described, increasing attention was given to their relationships and to the manner in which they may be grouped in a classification. Earlier authors (e.g., Friese in 1895, W. H. Ashmead in 1899) arbitrarily divided bees into those which are parasitic and those which are not. The resulting classifications were highly artificial for they separated such obvi- ously close relatives as Bonibus and Psithyrus. An entirely new and carefully considered classification of bees was proposed by Robertson, a Carlinville, Illinois, schoolteacher and botanist, in 1904. Rob- ertson observed that the seventh abdominal tergum of many female and the eighth of many male bees bears a flattened, bare, margined pygidial plate. He MICHENER: THE APOIDEA ^jj believed that the presence or absence of this plate was a primary character di- viding bees into two great natural groiips. In fact, he went so far as to suggest that the pygidialate and apygidialate bees might have arisen from pygidialate and apygidialate specoid wasps, respectively. This classification had many merits but unfortunately Robertson, working with a limited fauna, did not realize that the pygidial plate could be independently lost in various groups. Borner devised another classification in 1919, based primarily on mouth- parts. Like the other classifications which utilize chiefly one set of characters, this resulted in some artificial arrangements. A serious attempt to use all available characters was made by the present author in 1944. The result was a classification quite different from previous ones. It is to be hoped that as more characters are discovered and utilized, this clas- sification will be modified to refiect the added knowledge thus obtained. Bionomics Most students of bees have been interested in bionomics of these insects. That a considerable amount of information on this subject was available for European species a century ago is shown by leafing through Part I of F. Smith's Catalogue of British Hymenoptera in the Collection of the British Museum (1855). Subsequent work has mostly been done in Europe, with German work- ers taking the lead in work on social Halictidae, Italian workers (especially Guido Grandi) on larval characters, with individuals of all principal nations contributing papers on nest-making and habits of various groups. Outstanding among students in this field was Malysliev. In America work on wild-bee biology has lagged almost until the present time, although several workers are now in- terested in such studies. The importance of bees in cross pollination of various plants has been long understood, but only within the last fifteen years has the superiority of certain solitary forms over the honeybee for pollination of some crop plants been real- ized. This realization has provided a stimulus to the study of bionomics and several persons are now investigating these matters in the hope of solving prac- tical pollination problems. Some groups of bees of special interest for various reasons have received a great deal of attention. Outstanding, of course, is the honeybee, upon which much has been written. No discussion of this sort would be complete without mention of the famous studies by von Frisch, still under way, on the behavior and sense physiology of this insect. Another major group of social bees, the Meliponini, has received consideralile study, for example from von Ihring in Brazil and more recently from H. F. Schwarz in this country and Warwick E. Kerr in Brazil. The Future The lines of investigation which now seem important for further study among bees are numerous. For example it seems certain that among the Halic- tidae every gradation between solitary and thoroughly social forms will be found and that studies of this grading series will shed light on the steps in de- 578 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES velopment of social organization and the forces acting to cause such develop- ment. Further comparative studies will shed a flood of light on the evolution of instincts. Morphological studies of many sorts will provide further information on phylogeny, which is needed to verify or alter the present classification. Studies of such matters as parallelisms, orthogenesis, and the like can then be approached on a sounder basis. Biosystematic studies of all groups will add to our general knowledge of bee species, their ecologies, and their evolutionary and distributional patterns. In this connection a matter of special interest concerns pollen-collecting habits. Many species, termed polylectic, collect pollen from all sources; others, known as oligolectic, from only a few related species of plants. The evolution of this specialization, or the general problem of host specificity, can well be studied among bees for every intergrade between oligolecty and polylecty exists within numerous genera. These are merely some of the biological problems upon which bees may well provide information and upon which the present author and his students and associates hope to work. We trust that others will help, for there is work enough for many. Some will prefer to work on quite different problems, for example, pollination, sense physi- ology, and so forth. One of the great troubles with most entomological papers is that they are written to be read by only a very few specialists. They provide a mass of minutae and few generalizations. Let us hope that more and more entomologists will attack and solve, through the insect groups in which they specialize, problems of general biological interest. Too many gather the needed data and are con- tent to publish them without analysis, ignorant of or indifferent to the biologi- cal principles to which these data may contribute. DIPTERA Charles P. Alexander University of Massachusetts, Amherst As at present known, the Diptera or two-winged flies comprise the fourth largest order of insects, with approximately 85,000 described species, which pos- sibly is not more than some 20 per cent of the total number in existence. Some of the better known countries and states have species of Diptera about as fol- lows: Great Britain, 5,200; United States, 16,700; New York, 3,615; New Eng- land, 3,325; Michigan, 3,235. Various classifications of the order have been proposed, the most recent by Hennig (1948) which separates the Diptera into two suborders, Nematocera, with sections Bibiomorpha and Culicomorpha, and Brachycera, with sections Ta- banomorpha and Muscomorpha. A widely accepted arrangement, which is fol- lowed in this paper, divides the order into two suborders, the Orthorrhapha with two series, Nematocera and Brachycera, and the Cyclorrhapha with three series, the Aschiza, Schizophora, and Pupipara. In the following brief account, the leading events and many of the outstanding workei-s are indicated, together with ALEXANDER: DIPJERA 579 significant dates of publication. Complete references may be checked in Hagen's Bibliotheca Entomologica, The Zoological Record (1864-clate), and in other standard works. First Period, 1758-1853 Linnaeus (1758) recognized only ten genera of Diptera, with no distribu- tion into families, and including only 188 species. These genera in their exact arrangement and with the number of included species are as follows : Genus Species Genus Species 220. Oestrus 5 225. Empis 3 221. Tipula 37 226. Conops 6 222. Musca 100 227. Asilus 12 223. Tabanus 12 228. Bombylius 3 224. Culex 6 229. Hippobosca 4 Virtually all of the species were from Europe and chiefly from Sweden, with a very few from North America. That Linnaeus had no idea of systematic inter- relationships is shown by his separation of the two Nematocerous groups, Tipula and Culex. Linnaeus' outstanding entomological student, Fabricius, greatly in- creased the number of species, both from Europe and abroad, and in introduc- ing his so-called Cibarian system of insect orders, based on a study of their mouthparts, proposed the ordinal name Antliata to replace the Linnaean term Diptera, a suggestion that found little or no acceptance among later workers. In 1800 there appeared a highly controversial paper by Meigen, the "Father of Dipterology," followed (1803-1838) by a series of notable works by this same student. Toward the end of the period several workers appeared, including Mac- quart (1838-1855) and Wiedemann (1819; 1828-1830), whose principal publi- cations were on exotic Diptera, then becoming available in some numbers through various scientific expeditions. Other taxonomists included Curtis (1824-1840), Fallen (1814-1825), Haliday, Latreille, Robineau-Desvoidy (1830), Say, Schel- lenberg (1803), Stephens (1828-1846), and, toward the end of the period, West- wood (1839-1840), Zetterstedt, and Francis Walker. In dipterous morphology, important basic work was done by Latreille (1825), who proposed terms such as prothorax, mesothorax, and the like, and by Audouin (1824-1832) who further refined the terminology of the thorax, giving us such familiar terms as scutum, praescutum, scutellum, episternum, and many others. In biology and life histories, the early studies by Swammerdam were carried for- ward in the notable works of De Geer and Reaumur. The first general textbooks on entomology were prepared by Burmeister, Kirby and Spence, and Westwood. Second Period, 1853-1903 The close of the preceding period and the virtual end of the belief in "fixity of species" with the publication of Darwin's Origin of Species (1859), intro- duced a new and vigorous epoch. The pre-Darwinian belief resulted in an almost incredible synonymy in the order, as exemplified in an extreme instance in the posthumous work of Robineau-Desvoidy (1863), wherein the common parasitic fly, Tachina vulgaris Fallen, was redescribed no fewer than 245 times, the sup- posed species being distributed in five different genera! In 1853, museums and collections containing Diptera were generally small 580 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES and scattered. There was not one in the United States except the small Harris col- lection in Boston. In Europe, the leading collections were in London, Oxford, Paris, Leningrad, and Vienna. This neglect of the order was destined to be changed in an almost dramatic manner following the arrival in America in 1856 of Baron Osten Sacken, "Father of American Dipterology," who served as Sec- retary of the Russian Legation in Washington until 1862, and as Russian Con- sul General in New York from 1862 to 1871. Osten Sacken himself was one of the most accomplished students of the order, but served an even more important function in providing ample materials of North American Diptera for the study of Hermann Loew, outstanding systematic dipterologist of the period. Loew de- scribed as new some 1,350 species of North American Diptera, chiefly in a series of ten reports, or centuries (1861-1872), each including one hundred species. The combined Loew-Osten Sacken collections, now preserved in the Museum of Comparative Zoology in Cambridge, Massachusetts, comprise the most important basic series of flies in America. Other outstanding European students of Diptera who were most active dur- ing this period included Becker, Bellardi, Bergenstamm, Bergroth, Bigot, Bons- dorff, Brauer, Dziedzicki, Egger, Gerstaecker, Giglio-Tos, Girschner, Jaennicke, Karsch, Kowarz, Lioy, Meade, Mik, Pokorny, Portschinsky, von Roder, Rondani, Riibsaamen, Schiner, Schnabl, Stein, Strobl, Winnertz, van der Wulp, Zeller, Zetterstedt, and various others. The outstanding major works of this group in- cluded Brauer und Bergenstamm 's Die Ziveiflilgler, 7 parts (1880-1894), Ron- dani's Dipterologiae Italicae Prodromus, 8 volumes (1856-1880), Schiner's Fauna Austriaca, Diptera (1862-1864), and Zetterstedt's Diptera Scandinaviae, 14 volumes (1842-1860). In North America, in addition to the work of Loew and Osten Sacken, this period marked the initial activity of Coquillett, Johnson, and Williston. Lead- ing workers in South America included the Lynch Arribalzagas and Philippi (1865). In Australia, a most outstanding figure was Skuse, whose eight princi- pal papers on the Diptera of Australia (1888-1891) are of unusual importance. Virtually all other taxonomic work on exotic Diptera was accomplished by stu- dents in America and Europe, including Bellardi, Schiner, and van der Wulp. In the fields of dipterous morphology, phylogeny, and biology noteworthy advances were made. The science of chaetotaxy was proposed and developed by Osten Sacken (1881), although the term "machrochaeta" had been suggested many years before by Rondani (1845). A major landmark was attained in 1883 when Brauer first demonstrated the importance of the larva in classification and used the nature of emergence from the pupa to furnish the primary division of the order into Orthorrhapha and Cyclorrhapha. Weismann (1864) published an outstanding paper on dipterous development. Our state of knowledge of embryology was indicated by Korshelt and Heider (1890-1892). Pioneer work on the venation and morphology of the wing was accomplished by Adolph (1879), Amans (1885), Cholodkowsky (1886), and others. Loew and Schiner proposed their respective systems of venation in 1862. Significant work on the morphology of individual dipterous types was done on the blowfly by Hammond (1881) and Lowne (1890-1895). The earliest work on fossil Diptera began at this time with the appearance of Loew's paper on the Amber Diptera (1850). He was followed by several ALEXANDER: DIPTERA 581 other students including Brongniart (1878), Forster (1891), Giebel (1862), Heer (1849-1865), Heyden (1870), Meunier (1892-1917), Oustalet (1870), No- vak (1877), Scudder (1890-1894), and others. The first catalogues of Diptera, covering various regions appeared, including Osten Sacken (1858; 1878) for North America, van der Wulp (1896) for south- ern Asia, Reed (1888) for Chile, and others. An outstanding event of the period was the publication of Scudder 's NomencJator ZooJogicus (1882). Important general texts include those of Comstock and Packard, Third Period, 1903-1953 At the very end of the preceding period, the discovery that certain blood- sucking insects and other arthropods carried diseases of man and other animals, focused attention sharply on the various families of Diptera that might be in- volved, including the Psychodidae {Phlehotomus) , Ceratopogonidae {CuUcoi- des), Culicidae, Simuliidae, Tabanidae, and various muscoids, and including also, because of its habits, the housefly. There followed intensive work on all of these groups from every possible aspect. These initial studies led to the publi- cation of monographic works on mosquitoes by Theobald (1901-1910) and by Howard, Dyar and Knab (1912-1917), as well as a multitude of other papers and reports on the group, chiefly by Blanchard, Coquillett, Cristophers, Dyar, Giles, Goeldi, Graham, Peryassu, and others of the earlier period, and by Bar- raud. Bonne, Bonne-Wepster, Costa Lima, Edwards, Evans, Lang, Lutz, Mar- tini, Matheson, Newstead, Patton, Shannon, Taylor, Wesenberg-Lund, and others of the intermediate period. At a still later date, especially during and after the recent war, a host of younger students have almost completely revolutionized our knowledge of mosquitoes, particularly from the tropics. Similarly, in the other groups of blood-sucking flies above mentioned, many capable workers have advanced our knowledge far beyond that of most other groups of Diptera that are not of medical importance. It is a matter of regret that restrictions of space prevent the listing of such students. In the field of general dipterous taxonomy, the period likewise produced numerous workers. Some of these, particularly in the earlier years when the number of described species was still not excessive, were able to study certain families for the entire earth, while others were able to name many of the com- mon flies of a more restricted area. There remain only a few such broad students of the order and we definitely have entered a period when specialization seems required. Among those students who have descriljed species in both suborders of Dip- tera are the following: Abreu, Aldrich, d'Andretta, Austen, Pereira Barretto, Becker, Bezzi, Brunetti, Coquillett, Curran, Duda, Enderlein, Engel, Fairchild, Frey, Johnson, Knab, Lutz, Mackerras, Malloch, Matsumura, de Meijere, de Meillon, Pritchard, Seguy, Shannon, Shiraki, Stone, Strobl, Taylor, Verrall, Wil- liston, Wirth, and some others. Some of the leading workers on the taxonomy of the Nematocera include the following: Abonnenc, Alexander, Barnes, Borel, Brug, Causey, Damasceno, Dampf, Doane, Dyar, Felt, Floch, Fox, Freeman, Goetghebuer, Hertig, Hoff- man, Holmgren, Ingram, Johannsen, Kieffer, Kitakami, Komp, Lackschewitz, 582 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Landrock, Lane, Lee, Lengersdorf, Lundstrom, Maefie, Mangabeira, Mannheims, Martini, Natvig, Parrot, Pierre, Riedel, Eogers, Rozeboom, Sasa, Satchell, Shaw, Smart, Theodore, Tokiinaga, Tonnoir, Townes, Vargas, West, and many others. The chief students of the Braehyeera and Cyclorrhapha include among others : Aczel, Arias Encobet, Aubertin, Bau, Bequaert, Bromley, Brooks, Brues, Carrera, Cole, Collin, Cortes, Cresson, Czerny, Duda, Efflatoun, van Emden, Fer- guson, Ferris, Fluke, Goffe, Hall, Hallock, D. E. Hardy, G. H. Hardy, Hendel, Hennig, Hering, Hermann, Hesse, Hine, Huckett, Hull, James, Karl, Kertesz, Krober, Lichwardt, Lindner, Lundbeck, Melander, Metcalf, Miller, Munro, New- stead, Oldenberg, Oldroyd, Olsuf'ev, Pantel, Paramonow, Parent, Patterson, Patton, Philip, Pleske, Reinhard, Ricardo, Ringdahl, Sabrosky, Sack, Schmitz, Schuurmans-Steklioven, van Schuytbroeck, de Souza Lopes, Speiser, Stackel- berg, Stein, Steyskal, Stuardo, Surcouf, Szilady, Townsend, Villeneuve, Zia, and many more. Great progress was made in the study of dipterous morphology, biology (in- cluding genetics), and embryology. In morphology, outstanding w^ork was ac- complished by Crampton (1909-1943), Ferris and his students, and Snodgrass (1909-date). A detailed bibliography is provided by Crampton (1942). Studies of certain body regions include the head and mouthparts by Peterson, the ptili- num by Laing (1935), the thorax by Snodgrass and Young, the pretarsus by Holway, and virtually all structures of the body by Crampton. Detailed mor- phological studies of specific insect types include papers by Williams on the Tanyderidae, and by Bromley on the Tabanidae. In wing venation, the basic studies begun by Comstock and Needham at the close of the preceding period culminated in the major work by Comstock (1918). Modifications of the Com- stock-Needham system were proposed by Alexander, Bromley-Shannon, Goffe, G. H. Hardy, Lower, Seguy-Vignon, and others. Recent important texts have appeared covering the general subject by Berlese (1909-1925), Comstock, Imms (1925), Tillyard (1926), and others; on morphology by Snodgrass (1935) ; physiology by Wigglesworth (1939, 1950); and embryology by Hagan (1951) and Johannsen and Butt (1941). In biology, very numerous papers on the immature stages were published, these being summarized in full by Hennig, 3 volumes (1948-1952). Some of the more important works on dipterous biology included those of Alexander (1920) ; Chu (1949) ; Demerec, on DrosopMa (1950) ; Fabre (1913) ; Johannsen, on aquatics (1905-1937) ; Malloch (1917) ; Melin (1923) ; Miall, Peterson (1951) ; Phillips (1946); Rogers (1926-date); Thienemann, on aquatics (1914-1921); Usinger and LaRivers, on aquatics (1948); Wood (1952), and many others. In genetics, the importance of certain flies, especially Drosophila, and to a lesser extent Sciara, has produced an almost unparalleled amount of research by many students, including two Nobel prize winners, Morgan and Muller. Other leading workers are Bridges, Metz, Patterson, Sturtevant, Beadle, and others. Research on fossil Diptera was stimulated by the appearance of the major work by Handlirsch, Die Fossilen Insekten (1906-1908). Particular attention was devoted to the Florissant and the Baltic Amber (Bernstein-Forschungen, 1929-), by Alexander, Andree, Brues, Cockerell, Edwards, and others. Marked impetus was provided in the study of the order by the appearance of various catalogues, manuals, and faunal treatments. ALEXANDER: DIPTERA 583 Catalogues: The outstanding catalogue is Kertesz, Catalogus Dipterorum (1902-1910), the seven volumes covering the world fauna but being completed only to the Cyclorrhapha Schizophora. Other catalogues covering more restricted areas include Aldrich (1905) for North America; Becker, Bezzi, Kertesz, and Stein, 4 volumes (1903-1907), the Palearctic Diptera; Brunetti (1920), the Ori- ental region; Miller (1950), New Zealand; Wu (1940), China; Stuardo (1946), Chile. Still other catalogues treat individual families for limited areas. Lists: Of great value are the various local lists that indicate the extent of the fauna in any given area. Among such are the list of the British Insects, by Kloet and Hincks (1945) ; New England Diptera, by Johnson (1925) ; New York, by Leonard (1928); North Carolina, by Brimley (1938); and others. Genera Insectorum: This outstanding publication (1902-date) combines the systematic treatment to genera with a list of the world species. Several fascicles have appeared but the work is still incomplete, the authors of the published parts including Alexander, Bau, Brues, Edwards, Hendel, Johannsen, Keilin, Kellog, Kieffer, Krober, Melander, Pierre, Seguy, Surcouf, and Theobald. Faunal Treatments and Manuah: A large number of publications fall in the above broad classification. Manuals considering the North American fauna in- clude Williston (1908) and Curran (1934). Townsend's Manual of Myology, 12 volumes (1934-1942) considers the muscoidean genera of the world. Treatments for the major faunal regions include, for the Palearctic, Lind- ner's great work, Die Fliegen der palaearktischen Region, 8 volumes with nu- merous parts by many specialists (1923-date). Oldroyd, Freeman, van Emden, Smart, Collin, and others, the Diptera of the Handbooks of British Insects se- ries, volume 9 (1949-date). Seguy, Pierre, Goetghebuer, Kieffer, and others. Fauna de France, "Diptera" (1923-date). Lameer, Fauna de Belgique, "Dip- tera" (1907). Hendel, Hering, Karl, Sack, and others. Die Tierwelt Deutsch- lands, "Diptera" (1928-). Verrall, British Flies, "Syrphidae" (1901), "Stratio- myidae" (1908). Lundbeck, Diptera Damca, 7 volumes (1907-1927). Stackel- berg, higher flies of European Russia (1933). For North Africa and the Ethio- pian region, Efflatoun's Egyptian Diptera (1922-); Reports of the Ruwenzori Expedition, 1934-1935, published by the British Museum (1930-date) ; Explora- tion Pare National Albert, de Witte and Other Missions; Brussels (1937-date). For the Oriental region, the Fauna of British India, "Diptera," by Brunetti, Christophers, Barraud, Senior- White, Aubertin and Smart, 6 volumes (1912- 1941). In the New World, the Biologia Centrali- Americana, "Diptera," by Aldrich, Osten Sacken, Williston, and van der Wulp, 3 volumes (1886-1903). A most im- portant series of volumes on the Diptera of Patagonia and South Chile has completely revolutionized our knowledge of this critical region; 7 parts, several fascicles, by various authors (1929-1948). For northeastern North America, the important Diptera of Connecticut series, by various authors (1942-date). Other faunal treatments that may be mentioned include the Fauna Hawaii- ensis, Diptera, by Grimshaw (1901), and the Insects of Samoa, Diptera, by Col- lin, Edwards, Malloch, and others (1927-). Periodicals: Periodicals devoted entirely to the Diptera include the Encyclo- pedic Entomologique, "Diptera," Series B, by Seguy, Surcouf, and many others (1924-1940); those restricted to the order in large part are the Zeitschrift fiir 584 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES SystematiscJie Hymenopterologie und Bipterologie (1901-1908), Konowia (1922- 1931), and Insecutor Inscitiae Menstruus (1913-1926). Modern Control of Dipterous Pests: The Diptera are of economic importance chiefly through their attacking man and animals and by the transmission of various diseases, as discussed earlier. A further group of species destroy various crops, among such being certain gall midges, Cecidomyidae, as the hessian fly, pear midge, cloverseed midge, chrysanthemum midge, and others (Barnes, 1946- date) ; the fruit flies, notably the apple maggot, Mediterranean fruit fly, Mexi- can fruit fly, and many others; and a variety of pests that attack garden and forage crops, as the frit fly, cabbage maggot, and many others. Before the ad- vent of modern sj^nthetic insecticides (about 1945) a system of control had been established against most of these pests, based partly on chemicals, but also utiliz- ing biological and cultural methods. With the discovery of DDT and other well- known chemical compounds, very effective controls for many of these pests were obtained and it appeared that for certain of these, at least, the problem of hold- ing them in check had been solved. However, at the present moment it has become apparent that certain of these insects have built up a strong resistance to all such types of chemicals and it appears that it is only a question of time before we will have to revert, at least in part, to former methods of control. Such statements apply specifically to the housefly and mosquitoes but appar- ently it eventually will apply also to most if not all of the other forms against which such chemicals are now used. The Future The vast increase in our knowledge of the Diptera during the past century seems certain to continue in every field of study. As regards taxonomy, it is certain that far less than one half of the species in the order have been described and, as indicated previously, it seems very possible to me that perhaps only some 20 per cent may have been made known to this date. The airplane and other methods of modern transportation will enable collectors to visit the remote spots of the earth and the great museums will continue to grow apace. The value of the type specimen has become increasingly apparent and every possible precau- tion should be taken to safeguard such unusually valuable specimens against loss from fire, atomic destruction, or from any other cause. As an added pre- caution, wherever possible, such types should be photographed or so illustrated that there remains no possible question as to the identity of the species. As the number of described Diptera increases, students of the world fauna will of neces- sity be compelled to restrict their studies to individual families or perhaps even to lesser categories, such as genera. Already there are certain genera in the order with more than 1,500 described species, with many more awaiting discovery. Until very recently work on the taxonomy of any major group of insects was possible only lo students who were connected with leading museums or uni- versities that possessed unusually complete library facilities. The development of the microfilm and photostat processes, with other methods of reproducing literature quickly and economically, has changed this picture and it is now pos- sible to procure copies of papers in rare or otherwise virtually unobtainable HOLLAND: SIPHONAPTERA 585 publications by the microfilm process, thus enabling students to work while far removed from major libraries. As the species of any given region become better known, more attention will be devoted to the study of their biology and ecology. Compared to the number of described adults, only a small percentage of flies are known in their early stages and most of these are in groups of medical or economic importance. Simi- larly, under the impact of the so-called "New Systematics," increased attention will be devoted to a critical analysis of supposedly valid species in relation to clines and infraspecific categories. This analysis will result in a reduction in the number of supposed species but should be compensated for by the discovery of still unknown valid species. These are merely indications of some of the problems that must be consid- ered in the future. A fascinating field awaits the young entomologist who de- cides to devote his life and energies to a study of the Diptera. SIPHONAPTERA! George P. Holland^ Sustematic Entomology Unit, Division of Entomology, Ottawa, Canada The fleas constitute one of the smaller orders of holometabolous insects. About 1,350 species and subspecies, belonging to approximately 200 genera, are recognized at this time. In the adult stage, fleas are ectoparasites of mammals or birds. Their small size, the difficulty of collecting them (except for a few species!), and the lack of suitable techniques and equipment for preparing and examining specimens made them unattractive subjects for study a century or more ago. It is possible, too, that in those early times the sordid circumstances generally associated with fleas discouraged attention from potential students, who turned their talents to problems involving more aesthetic creatures. Ferris (1951) quotes Denny (1842) concerning lice, which were similarly regarded: "... the author has had to contend with repeated rebukes from his friends for entering upon the il- lustration of a tribe of insects whose very name was sufficient to create feelings of disgust." Certainly, by 1853, fewer than 30 specific names for fleas had been proposed, and of these only about 17 are now considered valid. Linnaeus recognized only two species of fleas, the so-called human flea, Pulex irritans, and the chigoe, P. (now Tiinga) penetrans. In the early nineteenth century, the familiar "domestic" species from European dogs, cats, rats, house mice, and chickens were described by Curtis, Bouche, Bosc d'Antie, Schonherr, and Schrank. A few species from endemic European moles, hedgehogs, bats, badgers, squirrels, and birds were named during this period also, as well as an 1. Contribution No. 3054, Division of Entomology, Science Service, Department of Agriculture, Ottawa, Canada. 2. Head, Systematic Entomology. 586 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES echidna flea from Australia, and a giant flea from northern Canada. By 1853, only two genera, Pulex and Ceratophyllus, were recognized and no general clas- sifications had been attempted. The first systematic account of the order was that of Kolenati (1863), who recognized eight genera. The more conservative Taschenberg recognized but five in his important work (1880), which was the standard reference on fleas at the end of the nineteenth century, when three outstanding students of the order made their appearance in the literature. These were Julius Wagner of Russia, N. Charles Rothschild of England, and Carl F. Baker of the United States. These men had a purely academic interest in the fleas, for in those days the role of these insects as vectors of plague and other diseases was not known. The effect of the attack by this trio, and by some lesser students, on the virtually untouched fauna during the next few years is well demonstrated by three world lists published by Baker over a ten-year period. In 1895 he listed but 35 species (actually, he missed a few), which he placed in three families and six genera. In 1904 he catalogued 134 species, and in 1905, as a result of "a most extraordi- nary activity among students of this group," supplemented this list by approxi- mately 120 additional names, arranging the whole into eight families. About this time, the association between fleas and the dreaded l)u1)onic plague was proved in India. There followed immediately a tremendous increase of in- terest in these insects, and the few specialists available found their services much in demand. Baker ceased work on fleas in 1905, but Rothschild and Wagner con- tinued to occupy leading positions. The former purchased specimens from col- lectors all over the world, and, in 1915, established a publication {Ectoparasites) that was devoted almost exclusively to papers on the taxonomy of fleas. Dampf of Germany published a number of papers that were particularly well illus- trated for their time. Oudemans, the great Dutch acarologist, published papers on flea phylogeny, in one of which (1909) was proposed a subordinal division that was followed for many years and has been discarded only recently. The most outstanding student of fleas, the former friend and colleague of Charles Rothschild, is Karl Jordan, whose work on the order extends over half a century. First assisting Rothschild (illustrating many of the early Rothschild papers), then publishing jointly until the latter's death in 1923, and since then continu- ing alone, Jordan has described more species and exerted more influence on the development of a natural classification of these insects than any other individual. His nearest competitor was Julius AVagner, who left Russia after the revolution of 1917 to live in Yugoslavia. Shortly before his death, Wagner sold his collec- tion to the Staatsmuseum in Hamburg and it is known that the larger portion of it perished when the museum was destroyed by bombing during World War II. Wagner described many genera and species and published a number of works on flea morphology as well as a catalogue of the Palearctic species and several papers on classification, of which the most important appeared in 1939. The framework of our knowledge of the fleas of the world is based largely on the works of Rothschild, Jordan, and Wagner. The first half of the twentieth century has been a period of species descrip- tion and discovery of new specific distinctions. In 1900, for instance, Rothschild drew, for the first time, attention to the taxonomic value of the terminal ab- dominal segments of the female, and to specific differences in the spermatheca. HOLLAND: SIPHONAPTERA 587 Though important theories on phylogeny have been published and classifica- tions have been proposed, no really satisfactory arrangement is yet available. The lack of agreement on relationships is well illustrated by the treatment of Anomiopsylhis and related genera, which in three major works on North Ameri- can fleas published between 1942 and 1947 appeared in three different families. Jordan, of all students of fleas the most experienced and best equipped to pro- pose a general classification, has not done so, except for a limited but nonetheless important contribution in Smart (1948, rev. ed.). This neglect was in part de- liberate, Dr. Jordan being reluctant to embark prematurely upon so difficult a proceeding when new and unusual material was turning up continually all over the world. Nevertheless, it was his intention to prepare a monograph of the fleas of the world, but this was prevented by World War II. However, all is not lost, and G. H. E. Hopkins and the Hon. Miriam Rothschild (daughter of Charles) are now preparing a catalogue of the Rothschild collection largely ac- cording to Dr. Jordan's views on the phylogeny of the group. All students of Siphonaptera eagerly await the appearance of this work, which should provide the most acceptable classification yet developed.* The flea fauna of many parts of the world is now fairly well known; that of North America is particularly thoroughly investigated, in part because of con- cern over sylvatic plague, which is the manifestation of Pasteurella pestis in wild mammals. There have been numerous short papers by various authors, and larger taxonomic works have been published by Ewing, I. Fox, Holland, Hub- bard, and Traub, and a catalogue of literature by Jellison and Good. The fleas of western Europe are fairly well known, and a group of siphonapterists, led by loff, have made extensive contributions to the knowledge of fleas in the U.S.S.R. Bedford, deMeillon, and Hopkins have published on African fleas, Liu on those of China, and Cunha, Pinto, Guimariles, and others have made contri- butions from the Neotropical region. Sharif of India has made important con- tributions to morphology as well as to taxonomy, and Traub and Smit are cur- rently publishing descriptions of fleas from many parts of the world. In 1946, two papers of the greatest value to flea students were published. These were Snodgrass's account of the skeletel anatomy of fleas, and da Costa Lima and Hathaway 's catalogue to the literature on the order up to 1944. F. G. A. M. Smit of the British Museum at Tring recently circulated a list of about sixty contemporary students of fleas. Less than a score of these are really active in flea systematics, and most work at the species level. It is to be hoped that a number of students will devote their efforts to considerations of evolution and phylogeny of these insects so that a firm classification may ulti- mately be achieved. It is now pretty well conceded that the fleas are divisible into two major groups, usually considered as superfamilies, the Pulicoidea and the Ceratophyl- loidea. These in turn may be divided into about 50 or more fairly well defined natural groups of genera, the arrangement and constitution of which are the basis for much present-day disagreement. Related species and genera may be as- sembled fairly conveniently, but a number of cases of wrong association through superficial resemblances brought about by adaptation remain to be sorted out. *The first volume (of five) of this work has been published (1953) since this manuscript was prepared. 588 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES But the relationship of the groups to each other and their arrangement as a series of families, or as a hierarchy of families, subfamilies, and tribes, poses more difficult problems. The origin of fleas, too, remains obscure. There is virtually no fossil record. DeGeer first recognized them as ordinally distinct from other insects. Various authors would associate the fleas with, or derive them from Coleoptera, Diptera, Mecoptera, Trichoptera, or Ilemiptera. Jordan (1950) presented a provocative paper at the Eighth International Congress of Entomology at Amsterdam, and proposed that a symposium on the origin of fleas be organized for the Ninth Congress. That their association with mammals is of long standing is indicated by the host-relations of some groups today : a special family of fleas on bats, for example, and the so-called helmet fleas, which appear to be associated with mar- supials in the Neotropical and Australasian regions. Many fleas exhibit a high degree of host specificity, and it is clear that many evolutionary lines have died out with groups of mammals that have become extinct. Some relict species sug- gest, in tantalizing fashion, some of these losses to the flea student. The primi- tive sewellel {ApJodontia rufa), besides having a parasitic beetle and two aber- rant species of mites, supports four species of fleas, three of which belong to monotypic genera (two of these genera might well be placed in special sub- families) and the fourth species is the largest of its genus and perhaps the world's largest flea ! The evolutionary picture is sketchy in the extreme and is complicated by numerous examples of convergence and host-transference, all of which make the study of flea phylogeny and host-relationship even more difficult. FOSSIL INSECTS F. M. Carpenter Harvat'd University Since students of insect paleontology are dependent on the discovery of insect-bearing deposits, progress in this field has lagged behind that of other aspects of systematic entomology. Investigations of a century ago were largely concerned with insects preserved in Baltic amber and the Solenhofen (litho- graphic) limestone in Bavaria, both of which had been known since the time of the Roman Empire. In 1853 the amber insect fauna was in the process of being described by G. C. Berendt (with the aid of Hagen and others), whose two-volume treatise (1845-1856) deserves to be ranked among the great classics on insects. Many Solenhofen insects had already been described by Germar (1842), who then (1853) turned his attention to Tertiary insects of Germany. The same year (1853), 0. Ileer published the last of his papers dealing with the Tertiary insects of Oeningen and Radoboj, the whole series of publications forming a volume of over six hundred pages. The Jurassic insects of England were being studied by J. 0. Westwood (1854) and Carboniferous insects from the Saar Basin by F. Goldenberg, who established (1854) the extinct order CARPENTER: FOSSIL INSECTS 589 Palaeodictyoptera. The first general survey of fossil insects was published at this time by C. G. Giebel (1856), Die Insekten und Spi7inen der Vorwelt; this was a systematic review of all known fossil insects. Shortly after this, in 1865, S. H. Scndder published the first of one hundred thirty papers which were to appear on fossil insects before his death in 1910. His contributions were by far the most important in the field. Most of his de- scriptive accounts dealt with North American material but his more general treatises were world-wide in scope. Included among the latter were his Classed and Annotated Bibliography of Fossil Insects (1890) ; Index to the Known Fos- sil Insects of the World (1891); and Systematic Review of Our Present Knowl- edge of Fossil Insects (1890). His Tertiary Insects of North America (1890) is on a par with Berendt's work on amber insects mentioned above. The discovery of insects in the Carboniferous shales of Commentry, France, in 1875, led to a notable contribution by C. Brongniart, Recherches pour servir a Vhistoire des insect es fossiles des temps primaires (1894), in which the first specimens of giant Protodonata were described. Shortly after the beginning of the present century, Handlirsch's Die Fos- silen Insekten appeared (1906-1908). This, another classic in entomological lit- erature, had a profound influence on the ordinal classification of insects in general. His Revision des palaeozoischen Insekten (1919) and the posthumously Neue Untersuchungen ilher die fossilen hisekten (1938-1939) were hardly more than superficial reviews of the literature. The chapter on insect paleontology which he contributed to Schroder's Handhuch der Entomologie (1921) and which contained many highly imaginative restorations, is his best known work on this subject. Although many other entomologists, in addition to Handlirsch, have pub- lished on fossil insects during the past fifty years, only four have made insect paleontology their major field of study. These are: T. D. A. Cockerell, who de- scribed a great many insects from Tertiary deposits in Colorado, belonging to nearly all orders; R. J. Tillyard, whose stimulating papers on Permian and Meso- zoic insects and on insect phylogeny in general aroused the interest of ento- mologists in these subjects: A. V. Martynov, whose investigations on Russian material have added enormously to our knowledge of Permian and Jurassic in- sects; and F. M. Carpenter, who has been chiefly concerned with the Carboni- ferous and Permian insects of North America and with the evolution of insects in general. In addition, mention should be made of A. Lameere, who, although he published only a few papers on the subject, made a significant contribution to general aspects of fossil insects. Many other entomologists, far too numerous to be mentioned here, have made significant contributions on the geological history of particular groups of insects. The following might be mentioned as examples only: W. M. Wheeler (ants), C. T. Brues (parasitic Hymenoptera and phorid flies), F. Meunier (Dip- tera), G. Ulmer (Trichoptera), C. P. Alexander (Tipulidae), H. F. Wickham (Coleoptera), F. M. Hull (syrphid flies), F. E. Zeuner (Orthoptera), G. Statz (Diptera), E. E. Bekker-Migdisova (Hemiptera), and B. B. Rohdendorf (Dip- tera). In addition, a number of paleontologists have dealt with the insects of certain formations, notably II. Bolton (Carboniferous of England), P. Pruvost (Carboniferous of Belgium), and P. Guthorl (Carboniferous of Germany). 590 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES At present our knowledge of Tertiary insects exceeds that of any other past geological period; that of the Permian is the next best known. Most needed, therefore, are collections from other periods, especially the Cretaceous, which is almost a blank, so far as insects are concerned. Eevisional studies of previously described material by specialists in certain orders are also needed, as well as investigations on unworked material. The most extensive collection of fossil insects, comprising about 60,000 specimens (including the Scudder Collection), is contained in the Museum of Comparative Zoology. Other important collections are in the British Museum (Natural History), the Institute of Palaeontology in Moscow, the Museum National d'Histoire Naturelle, in Paris, and the United States National Museum. The collection of amber insects, formerly housed in Albertus University at Konigsberg and including about 100,000 specimens, was destroyed during the Second World War. HERPETOLOGY Bij KAEL P. SCHMIDT Chicago Natural History Museum Expanding herpetology/ like a branching tree, underwent development in various directions in its new growth during the latter half of the last century and the first half of the present one, and even within any one of these branches there may be varied directions of interest that require some disentanglement. In the present historical essay I have not attempted any unified arrangement, but have followed the branches or the individual twigs of the tree of herpetology as they have seemed important or interesting. Herpetology may be broadly interpreted as including every phase of biological studies in which identifiable species or higher groups of amphibians and reptiles appear, and is so interpreted here. Emphasis, however, is upon the history of description and classification of the existing world fauna, which involves the story of the exploration of the world for the several thousand species of amphibians and i-eptiles. Most of the rise of our knowledge of the extinct members of these two groups falls within the century 1850-1950, but this segment of our history cannot be elaborated in the present essay. Emphasis on the field of systematics, the central trunk of our tree, carries with it an interest in the natural histoiy of the amphibians and reptiles. Natural history I interpret as the less critical forerunner of a more critical science of ecology. Even without this modern development, the natural history of the crea- tures in question has the merit of affording a base for the popular and semipopular literature of herpetlogy, which brings its more seriously scientific studies into the domain of knowledge of the general public and gives school children a key to a segment of the zoological sciences. This department of herpetological literature is peculiarly rich and requires some attention in a historical review. In addition to systematics, geography, and general natural history, the principal develop- ments in anatomy, physiology, embryology, and behavior are of major importance to a broad view of the liistory of herpetology. Finally, within each of the separate fields, historic interest focuses upon the personalities of the individuals who initiated fruitful directions of investigation or dominated them. The principal museums of the world have had pre-eminent roles in the growth of systematic studies and in the exploration of the world for new species. Thus the hierarchies and successions of the museum herpetologists become important. The fact that 1. studies on amphibians are commonly combined with those on reptiles in the zoo- logical subscience "Herpetology." These creatures compose respectively the class Am- phibia and the class Reptilia, two of the major groups of backboned animals, which were long combined in the Linnaean class Amphibia. The animals in question, the salamanders, frogs, and caecilians (the living amphibians as now understood), and the turtles, croco- dilians, lizards, snakes, and the tuatara (the existing reptiles), were all commonly lumped together as reptiles in the popular mind and, for that matter, still are. The zoological distinction between the Amphibia and the Reptilia, though fully established, had not yet been properly carried through in general works at the middle of the nineteenth century. [591] 592 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES the museums introduce an element of nationalism, sometimes of nationalist riv- alry, adds interest to the story. The Era of Dumeril and Bibron A major summary of the field of herpetology, in ten volumes, marks the end of the first half of the nineteenth century. This is tlie Erpetologie generale ou Mstoire naturelle complete des reptiles, by Andre Marie Constant Dumeril (b. 1764, d. 1860) and Gabriel Bibron (b. 1806, d. 1848), based largely on the collections accumulated at the J\Iuseum of Natural History in Paris. The first volume of this work appeared in 1934, and the last in 1854; after the death of Bibron in 1848, A. H. A. Dumeril, the son of the senior author, aided with vol- umes 7 and 9. The tenth volume is an atlas of 120 colored plates. This work, still much referred to, gives a comprehensive scientific account of the reptiles in general (including the amphibians as the distinct order Batrachia), as to their structure and physiology as well as their systematics, together with an historical account of the literature of the subject, and this is supplemented by a general account for each of the principal orders recognized. One hundred and twenty- one species of turtles, 468 lizards (with which are included the crocodilians), 586 snakes, and 218 "batrachians" are described. The classification of the snakes foreshadows the more modern ones of Cope and Boulenger in being based on dentition; five equivalent groups are recognized, the Opoterodontes, the Aglypho- dontes, the Opisthoglyphes, the Proteroglyphes, and the Solenoglyphes. The last four terms were to become current herpetological property, useful even when their systematic importance was seen to be less than at first thought. The work greatly multiplied the number of known families of snakes, recognizing no less than twenty for the nonvenomous forms. The Erpetologie generale was the crowning work of a century of herpeto- logical studies, during most of which the leadership in the field had lain with the French. Earlier comprehensive treatments of the amphibians and reptiles had been supplied by the various editions of the Histoire naturelle of Buffon and the Regne animal of Cuvier. As a summary of what was known of the herpetology of the world in 1850, the Erpetologie remains a work of major importance. A direct line of succession of herpetologists at the Museum National d'Histoire Naturelle at Paris carries on from Constant and Auguste Dumeril through Leon Vaillant, P. Mocquard, and Fernand Angel (who died in 1950), to Jean Guibe. The most notable achievement of these generations was the herpetological explora- tion and description of the French colonies, especially of the great and remark- able island of Madagascar. The Dumerils were not left unaided at the National Museum in Paris after the death of Bibron. Marie-Firmin Boeourt, who came to the museum as pre- parateur in 1834 at the age of fifteen, became a competent herpetological artist as well as field collector. His first expedition was to Siam in 1861-1862; in 1864 he was placed in charge of the Mission Scientifique au Mexique et dans I'Amerique Central, an adjunct to the attempt of Napoleon III to establish a Mexican empire under the ill-fated Maximilian. The failure of the Mexican venture sent Boeourt to Guatemala and other parts of Central America. After his return in 1867 he devoted himself to the report on his collections of reptiles, and more especially SCHMIDT: HERPETOLOGY 593 ERPfiTOLOGIE GfiNERALE 00 HISTOIRE NATURELLE COMPLETE DES REPTILES, Par A.-M.-C. DUMERIL , MGMBnB DB l/lNSTITOT , PROFESSEUR PR I.A FACULTE DH MEDECIITS, PROFBSSEUR ET ADMINISTRATEUR DU MUSF.UM d'hISTOIRE NATURELLE , BTC. EN COLLABORATION AVEC SES AIDES NATURALISTES AU MUSEUM , FEU G. BIBRON , PROFESSEUR D'hISTOIRE NATURELLE A l'eCOLR rRUIAlRE SUPKRIBURE D8 tA VlUt DE PARIS ; ET A. DUMERIL . PROFESSEtlR ACRKGlfi DE LA FACULTE DK JU'UECINE POUR l'aXATOMIE ET LA PHTSIOtOOlt. ATLAS RENFERMANT 120 PLANCHES GRAVIES SUH ACIEB. PARIS. UBRAXaZi: EMCYCLOFEDZQirC DS HORETf RUB lUUTEFRUILLB , 12. 1834. Figure 1. Title page of ttie Erpetologie gin6rale. 594 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES to the production of 95 of the 101 plates in the accompanying atlas. Since he illustrated many of the type specimens from other collections and other museums, this great work remains one of the basic sources for studies on the Central Amer- ican region. The amphibian section, with twenty-one plates, is by Paul Brocchi, and the reptile section was completed by Mocquard, the last six plates being by Fernand Angel. Another direction of interest in herpetological studies in France may be seen in the continued attention to the fauna of France itself. As this became well explored as to systematics and geographic distribution, there arose opportunity for detailed attention to problems of life history and behavior. Leaders in this important reorientation of interests were Fernand Lataste, most important in herpetological history as the mentor of G. A. Boulenger, and Raymond Rollinat, who will be remembered for his fifty-year-long interest in La vie des reptiles (1934). Partly as a result of the long history of technical herpetological studies in France, the popular and semipopular literature of herpetology in the French language is particularly rich. In the decades following the appearance of the Erpetologie general the pres- tige of leadership in systematic herpetology passed from Paris to London and Berlin. At Berlin the scientific productivity of Wilhelm Carl Hartwig Peters (b. 1815, d. 1883) spanned three decades of active publication during his regime as Professor of Zoology at the University of Berlin and Director of the Zoological Museum. His career began with an important personal zoological expedition to Africa, the Reise nach Mossamhique, which extended from 1842 to 1848. Like most of his contemporaries, he was equally interested in various groups of ani- mals, often combining the descriptions of new species of mammals and amphib- ians, or of snakes and fishes, and describing collections from three or four of the continents in the same paper. After his. return from Africa a steady stream of short papers, mostly descriptions of new species, came from his pen in every year until his last. The first of the five great folio volumes of the reports on the Beise nach Mossamhique appeared in 1852, the last in 1882. Of these the volumes on mammals, fishes, and amphibians- were by Peters himself. Great collections of fishes, amphibians, and reptiles were meanwhile accumu- lating at the museum of natural history in Vienna, where the leadership in ichthy- ology and herpetology had fallen to Franz Steindachner (b. 1834, d. 1919). Steindachner, though more eminent in ichthyology, founded an Austrian school of herpetologists. He joined the staff of the Naturhistorisches Museum in 1860, and his publications in herpetology continue from 1862 to 1917. His papers include, with numerous short notes and descriptions, the reports on the collec- tions of the Austrian Novara Expedition (1867) , and quarto papers on collections made by himself in Africa, southwestern Asia, Brazil, and the Galapagos Islands. Before we return to the main thread of the development of herpetological studies (in London), other direct derivatives of the Paris school may be men- tioned. The Russian Imperial Academy of Sciences, and the Museum of Natural History in St. Petersburg became centers of herpetological publication under the regime of Alexander Strauch (b. 1832, d. 1893). Strauch was born in St. 2. Boulenger remarks that Peters was the last important herpetologist to employ the Linnaean class Amphibia in its comprehensive sense, to include both amphibians and reptiles. SCHMIDT: HERPETOLOGY 595 Petersburg, came to the Zoological Museum of the Imperial Academy of Sciences in 1861, and became its director in 1879. His papers (all in German) in the Memoirs and Bulletin of the Academy (1862-1892) included revisions of the crocodilians, turtles, and viperine snakes of the world. Strauch's successor in herpetological studies was A. M. Nikolsky, whose first paper appeared in 1886, with comprehensive accounts (in Russian) of the amphibians and reptiles of the Russian Empire in 1915-1918 {Faune de la Russie). Jacques de Bedriaga (Rus- sianized to Yakov Vladimirovitch Bedryagha; born in 1854, publishing career 1874-1912) interested himself especially in the herpetology of the JMediterranean region, of Europe generally, and at last of Mongolia. His account of the frogs and salamanders of Europe, Die lurch fauna Europas (1889-1897), is a com- prehensive treatment of the fauna, though it suffers by comparison with Boulen- ger's magnificently illustrated work on the Tailless Batrachians of the same region. Bedriaga 's reports on the amphibians and reptiles of the Przewalski Expeditions to Central Asia amount to more than seven hundred pages (with parallel Russian and German text), and ten plates. In Italy the wealth of lacertid lizards, whose suitability for pets in terraria has always been a source of herpetological interest in Europe, and the somewhat richer Mediterranean fauna in general, gave rise to an early and continuing interest in herpetology, and to one of the earliest elaborate accounts of a regional fauna. The "Amfibi" (both amphibians and reptiles) constituting C. L. Bona- parte's Volume II of his Iconografia della Fauna Italica . . . , (1832-1841), con- temporary with the early volumes of the Erpetologie generate, depicted the am- phibians and reptiles of Italy on 53 colored plates. The review of the Italian herpetological fauna was redone by Lorenzo Camerano between 1883 and 1891. The tradition of such national faunal works continues to the present day. The director of the Museum of Natural History in Milan, Georg Jan (b. 1791, d. 1866), undertook the ambitious project of illustrating the snakes of the world. The coverage of this work was unhappily reduced by the refusal of the British Museum to lend its specimens to be drawn; but the 300 plates drawn and litho- graphed in uniform style by Ferdinand Sordelli (who completed the work after Jan's death) remain one of the monumental contributions to the illustration of the snakes of the world. The Iconographie generale des opkidiens was published in 50 livraisons, each with six plates (1860-1881). Opportunities for zoological exploration, often with governmental support, were presented in the foreign colonies of European nations, and these may domi- nate the herpetological interests of a national group. Such colonial exploitation is exemplified in the contributions of J. V. Barboza clu Bocage (b. 1823, d. 1895), Director of the Portuguese National Museum in Lisbon, whose publishing career began in 1864 and was climaxed by his volume H erpetologie d' Angola et du Congo, published in the last year of his life. The Era of GiJNTHER and Boulenger Reserving other developments in herpetology in various parts of the world and the early history of the field in the United States for later sections, we must turn to the dramatic transfer of leadership in the study of amphibians and rep- tiles from the Continent to Great Britain, and in particular to the British Museum 596 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES G. A. BOULENGER A. M. C. DUMeRIL |r GABRIEL BIBRON E. D. COPE SCHMIDT: HERPETOLOGY 597 in London. In retrospect, the major turning point is discernible in the appoint- ment of George Albert Boulenger to the curatorship of reptiles at the British Museum in 1880. Studies on the amphibians and reptiles preserved in the British Museum had been greatly promoted by the voluminous but uncritical work of John Edward Gray (b. 1800, d. 1875), who began the tradition of published catalogues of the museum collections. Gray's publishing career (1825-1874) marks the rise in importance of two journals that became the principal media for the description of new forms — the Annals and Magazine of Natural History and the Proceedings of the Zoological Society of London. Much of the most notable contribution made by Gray to herpetology was the choice of Albert Giinther^ (b. 1830, d. 1914) as his assistant in the divisions of ichthyology and herpetology, and, as it turned out, his successor as Keeper of Zoology. The young German (born at Esslingen, Wiirttemberg), after taking holy orders in 1851 in the Lutheran Church, was diverted into a zoological career by the lectures of Professor von Rapp at the University of Tiibingen. He took his degree as M.D. at that university in 1857, having meanwhile studied with the great anatomist Johannes Miiller at Berlin and with Franz Hermann Troschel at Bonn, served at St. Bartholomew's Hospital in London, and written a book on medical zoology (published in 1858). In 1857 he accepted an assistantship offered by Gray at the British Museum, in which he was to catalogue the fishes, amphib- ians, and reptiles. By 1859 the great Catalogue of Fishes was under way, and the catalogues of Batrachia Salienta and of Colubrine Snakes were both published in 1858. His largest herpetological work was the folio Reptiles of British India (1864) , published by the Ray Society. Giinther's most notable herpetological discovery was that the New Zealand tuatara is not a lizard but a living representative of an otherwise extinct order of reptiles, the Rhynchocephalia (1867). His contributions to ichthyology so much overshadow his herpetological work, that we tend to underestimate him as a herpetologist ; but his greatest contribution to herpetology was, in his turn, his choice of successor, which fell to a young Belgian, George Albert Boulenger (b. 1858, d. 1937). Before turning to the work of Boulenger and the Boulengerian era, it is necessary to note the work of the Biologia Centrali- Americana, and of John Anderson, the origins of which fall in the time of Giinther. The herpetological share in the Biologia Centrali- Americana was important to the growth of the British Museum collections and affords an example, on a grand scale, of the effective aid of amateurs to museum work. The history of the Biologia is an extraordinarily pleasant story of a friendship between two Cambridge University students in the eighteen-fifties. Osbert Salvin and Frederick Ducane Godman were drawn together by a common interest in natural history, and their com- panionship led from wild-fowling in the Cambridge fens to the biological explora- tion of a quarter of a continent, resulting in the magnificent monument of the 63 quarto volumes of the Biologia. The volume on amphibians and reptiles (1885- 1902), illustrated with 76 lithographic plates by the fine artists of the era, was prepared by Albert Giinther. Another notable herpetological career was that of John Anderson (b. 1833, 3. His full name, Albert Charles Lewis Gotthilf Giinther, was usually so shortened. 598 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES d. 1900). With a medical degree from the University of Edinburgh, Anderson went to Calcutta in 1864. His arrival was fortunately timed, for the collections of the Asiatic Society of Bengal were then being turned over to the government of India. A new museum building was to be erected, and John Anderson was named Curator in 1865 and Superintendent a few years later. He retired in 1886, to live in London, and spent his winters in Egypt. While in India, Anderson took part in the two Yunnan expeditions, whose zoological results appeared in 1878-1879 in two quarto volinnes, herpetologically important for their descrip- tion of the remarkable turtle fauna of southeastern Asia. John Anderson's career and interests fall sharply into an Indian and an Egyptian period. After his retirement he devoted hilmself (and no small part of his fortune) to the preparation and publication of the Zoology of Egypt. The quarto volume on amphibians and reptiles in this work (1898) is not only mag- nificent in format and illustration, but is one of the most competent and soundly useful of faunal works in the history of herpetology. George Albert Boulenger, born at Brussels in 1858, exhibited a passion for natural history at an early age, and specifically for the study of amphibians and reptiles. During his student days at the University of Brussels he engaged in the identification of the materials in the Museum of Natural History (the Belgian National Museum) and came under the influence of M. Fernand Lataste, whom he addressed throughout his career with affectionate regard. His first paper, pub- lished at the age of nineteen, a revision of the iguanid genus Laemanctus with the description of a new species from the collections of the Brussels Museum, is already in such competent and scholarly form that it might have appeared forty years later, when he was the acknowledged dean of European herpetologists. He was made assistant at the museum in 1880, but very soon resigned, on the invi- tation of Dr. Giinther to come to the British Museum to undertake a new edition of the catalogues of amphibians and reptiles, quite as Giinther himself had been invited by Gray twenty-two years earlier. It is easy to see that it was the favor- able impression made by the twenty papers published as the result of his work at Brussels that caused the young Boulenger to be invited to the most distin- guished herpetological position in the world. At the British Aluseum Boulenger immediately plunged into the work of revision of the classification of the amphibians, applying to the frogs and toads the system suggested by Cope in 1865, and literally bringing order out of chaos in this group. The volumes for Batrachia Gradientia (the salamanders) and Batrachia Salientia (the frogs and toads) appeared in 1882. Next came the three volumes for the Lizards, 1885-1887; the volume for Chelonians, Rhynchocephal- ians, and Crocodiles in 1889, and the three volumes for the Snakes in 1893-1896. Important contributions to the family classification were made throughout, and from the first, descriptions of the species not in the British Museum collections were included, so that these nine volumes constituted a summary of the world fauna for the classes Amphibia and Reptilia to the year 1896. Though his clas- sification of the amphibians has required complete revision, his arrangement of the families of reptiles is essentially that current in 1950. Concurrently with the great series of catalogues, Boulenger published no less than 279 herpetological papers in scientific journals in the sixteen years from 1881 to 1896, in addition to a volume on the amphibians and reptiles of British India. SCHMIDT: HERPETOLOGY 599 After completion of the catalogues, Boulenger continued with the descriptions of new species, reports on additions to the collection, and reports on individual collections from all parts of the world. His separately published subsequent works were the finely illustrated Tailless Batrachians of Europe, which reflects his principal contact with living amphibians and reptiles and his early interests in field observation (1896-1897); the compact little summary Les Batraciens et principal ement ceux d'Europe (1910) ; the "Reptilia and Batrachia" in the Ver- tehrate Fauna of the Malay Peninsula (1912); the Snakes of Europe with its admirable introduction on snakes in general (1913) ; and the Monograph of the Lacertidae (1920-1921) . Work on fishes in the British Museum began in 1887, and Boulenger thereafter continued to puljlish in both ichthyology and herpetology, with main interest on herpetology, much as Giinther had worked in both fields, with emphasis of ichthyology. His total list of publications in scientific journals amounted to more than 875, of which 618 were on herpetological and 257 on ichthyological subjects. This is ivitJiout enumeration of his more popular papers in The Field, Cou7itry Life, etc. This large list of publications reflects Boulenger's habit of rapid work, made possible by his having done the catalogue volumes, but this contained the seed of a weakness. His memory was phenomenal, so much so that he so readily recognized species that he had seen before that he was disin- clined to check identifications made' "through the glass"; and so great was his prestige among his colleagues that they also did not usually check his identifi- cations further. "When Clifford Pope and I were making a round of museum visits together in Europe in 1932, we could not help being amused at the dismay of some of our herpetological hosts when we questioned the determinations made by Boulenger on some casual previous visit, and insisted in our unbelieving way on having the jars opened so that we could examine the specimens more critically. Boulenger was not inclined to revise the keys for identification drawn up for the catalogues, and when these led him astray he sometimes described new species instead of making the revisions of his concepts that were indicated. For all of Boulenger's mastery of the world fauna, he displayed little under- standing of geographic distribution, and never alternated collecting and field studies with his work on preserved material in the museum. In still another respect his work was superficial — during the sixteen years of the production of the catalogue it was inevitably focused at the species level, and he displayed neither interest in nor understanding of the partition of species into subspecies, which has from the beginning, and of necessity, been based on more accurate knowledge of geographical and ecological relations. By no means an anti-evolu- tionist, the theory of evolution made astonishingly little impact on his thinking. The great series of catalogues appeared before the organization of the Inter- national Commission for Zoological Nomenclature. Having already chosen those names that seemed best to him from a welter of early synon>Tny, it is perhaps scarcely surprising that Boulenger sliould have been casually indifferent to the new rules and codes. It is easy to understand also, how annoying this indiffer- ence was to those who, like Leonard Stejneger, took the new attempt to codify and regularize zoological nomenclature so seriously that they could quarrel in print over the omission or addition of an I, or over an elaborately complex method of determining the type species of a genus. One of the most important accomplishments of the British Museum group was 600 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES 0 b /'.^//^/t a f/i f>iifj n m^niha d Figure 2. Specimen plates from Boulenger's Catalogues: a. From the Batrachia gradientia, 1882, 6. From the Batracha salientia, 1882. c. From the Lizards, 1885. d. From the Chelonians, 1889. These show (in modern halftone reproduction) the kind of litho- graphic engraving that characterized all zoological illustration up to 1900. SCHMIDT: HERPETOLOGY ^Q] the establishment of order in the rapidly expanding literature of zoology by means of the Zoological Record. This was founded by Giinther in 1864; Giinther him- self contributed the sections for Amphibia and Reptilia from 1864 to 1872; from 1873 to 1879 they were done by A. W. E. O'Shaughnessy; and Boulenger took this field over from 1880 to 1914. Boulenger retired from the British Museum and from herpetological studies on completion of forty years of service at the Museum, and returned to Belgium and to an early interest in the European wild roses. This was in 1920, and he lived for the seventeen years until his death in 1937 with scarcely a thought of herpetology — at any rate, with only three small additions to his list of papers. It is easy to point to the defects of Boulenger's old-fashioned taxonomic work in herpetology and to sympathize with Stejneger, who was not only exasperated at Boulenger's lack of interest in nomenclatural changes, but was quite legiti- mately critical of the superficiality and carelessness of his work. There was to Boulenger's credit, however, the fundamental reform of the classification of the two great classes of vertebrates, which is so much now taken for granted that we tend to forget its importance; and there was the "merit of his defects," the fact that he did accomplish a complete review of the two great classes at the species level in the phenomenally short time of sixteen years ; the two generations of his successors throughout the world have bogged down in the reviews of single families, and often enough made a life work of a genus. Systematic zoology needs its Boulengers. The expansion of systematic herpetology from Dumeril and Bibron to Bou- lenger, and to the present day, may be reflected in the numbers of living species known : Dumeril and Bibron, 1851f Caecilians 8 Salamanders 58 Frogs 152 Turtles 121 Crocodillans 14 Lizards 454 Snakes 586 The tuatara Lithographic Illustration of Amphibians and Reptiles Great contributions were made to the illustration of the amphibians and rep- tiles of the world during the last half of the nineteenth eentur3^ This was the era of lithography. Able artists who had the patience to draw the scale detail of reptiles and the extremely skilled engravers on stone produced an extraordinary series of illustrations so accurate that they have not been surpassed, and need only the modern supplement of photographs from life. The art and technique of lithography flourished throughout Europe. Among the artists available to Steindachner in Vienna Eduard Konopicky deserves especial mention for his ability to catch lifelike attitudes in lizards, as well as for the accuracy of his scale detail, which, it was said, made it unnecessary to examine the specimen. In England the expanding publications of the Zoological Society of London were richly illustrated with black and white and with colored mlenger Estimate, 1896 1950 43 70 130 240 1146 2200 219 265 23 23 1969 3140 1639 2530 1 1 602 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES lithographs. The great English herpetologieal artist of this era was G. H. Ford, who illustrated the numerous papers of Gray and Giinther, including the Reptiles of British India. Of the 216 plates that illustrate Boulenger's catalogues, 78 are by P. Smit, 68 by E. Mintern, 42 by Mintern and Green, 19 by J. Green, 8 by Edward Wilson, and 1 by H. Gronvold. Smit's work illustrated the papers in the Pro- ceedings of the Zoological Society of London between 1884 and 1900, and he produced forty of the fifty magnificent quarto plates in Anderson's Zoology of Egypt, the remaining ten being by H. Gronvold and J. Green. Green's colored lithographs continued to appear in the Proceedings in the first decade of the twentieth century, the last one in 1917. During the second decade of this century, lithographic illustration was superseded by the various photographic processes. The illustrations of the lithographic era had a curiously pleasing quality, in which scale detail was combined with shading, and the loss of this technique is a loss to zoological illustration. The illustrators of that period deserve a more extended essay in appreciation of their services to science. The Boulengerian Era in Europe The immediate usefulness of Boulenger's catalogues for putting in order the collections accumulated and accumulating in other museums, and the example of his numerous short fauna! lists, fixed the style of herpetologieal publication for two generations in Europe. This Boulengerian era on the Continent continued the heriDCtoIogical exploitation of the colonial empires, notably of the Netherlands Indies by a Dutch school still in the shadow of the great Hermann Schlegel; of the Belgian Congo by Boulenger's successors in Brussels,^ and of the German African colonies by a Berlin group. "^ None of these rose above an unthinking multiplication of morphological species. The synopses of the Amphibia Salientia in Bas Tierreich by Nieden (1923) and Ahl (1931) do not rise above this level. Typical of such active national herpetologieal exploitation of colonies is the somewhat later work of Guiseppe Scortecci, in Milan, and later in Genoa, on the expanding Italian colonial empire. His earliest papers (1928) reflect this interest, and one even suspects unrealized colonial ambitions in papers on the reptiles of Yemen. The Italian contemporary of Boulenger was Count Maria Giacinto Peracca (b. 1861, d. 1923), whose ample means enabled him to keep a terrarium on the scale of a large conservatory, in which Galapagos turtles wandered at will. His publishing career and association with the zoological museum of Turin ex- tended from 1886 to 1917. His interests were wide, with a long series of papers on South American herpetology. The Boulengerian era in Vienna included the colleagues and successors to Franz Steindachner. Friedrich Siebenrock made notable contribution to the anatomy and systematics of turtles (publishing career, 1892-1924). Otto Wett- stein (son of the eminent botanist) will be most remembered for his detailed account of the anatomy of the tuatara in the Kiikenthal HandhucJi der Zoologie (1931) . On the retirement of Wettstein, the division of herpetology at Vienna was 4. J. K. de Jong, Nelly de Roolj, P. N. Van Kampen, and L. D. Brongersma. Properly- representative of more modern ecological field observation, Felix Kopstein (-1940) may be named with this group. 5. G. F. de Witte and Raymond Laurent. 6. Gustav Tornier, Fritz Nieden, Richard Sternfeld, Ernst Ahl, and Giinther Hecht. SCHMIDT: HERPETOLOCY 603 placed in charge of Joseph Eiselt. Franz Werner (b. 1867, d. 1939), long active as a teacher at the University of Vienna, was, perhaps unfortunately, persona non grata at the Natural Ilistorj^ Museum in Vienna under the regime of Steindach- ner. Much of Werner's work is competent herpetology in imitation of Boulenger; his reputation is marred by a few papers in which well-known exotic snakes are described as new species and new genera. These are wholly incongruous with his technically competent general account of the Amphibia and the special treatment of the Apoda in the Handbuch der Zoologie (1930). His major contribution to herpetology is his two-volume account of the amphibians and reptiles of the world in the fourth edition of Brehm's TicrJehen (1912-1913) , which is of broad interest to zoologists in general. An independent herpetological center grew up at the Hungarian National Museum at Budapest under the influence of Lajos von Mehely (whose herpeto- logical publications begin in 1890) and Baron G. J. von Fejervary (first paper in 1910) . Mehely was naturally enough interested in the European herpetological fauna. For the lizards of the genus Lacerta, his ideas as to which were the primi- tive and which the derived forms differed sharply from those of Boulenger. It is somewhat surprising to find an extensive series of papers by this author on New Guinean and South American frogs. Baron Fejerviiry, who was succeeded by his wife as curator of the herpetological collections, is known for his studies of the fossil varanid lizards and their relatives. Most of these papers are in German, some in both Hungarian and German. Since World War II Mrs. Fejevary has been publishing in Hungarian without benefit of summary in another language. There has been active interest in herpetology in the several Scandinavian countries since the times of Linnaeus. This is reflected not only in active work on the North European fauna, but in an interest in the amphibians and reptiles from foreign countries. Collections from individual travelers and from expedi- tions have accumulated in the museums and university collections of Sweden, Norway, and Denmark, throughout the Boulengerian and post-Boulengerian eras and have formed the basis for numerous reports^ (mostly in English or German). Oskar Boettger (b. 1833, d. 1910), equally known for studies in malacology and herpetology, made the Senckenberg Museum at Frankfort on the Main a cen- ter of herpetological studies. His papers reflect an influence quite different from the direct colonial interest of the national museums, and one characteristic of Frankfort, for the numerous correspondents who sent him specimens were busi- nessmen with amateur interests in natural history, wlio took time to collect for him, and for the home town museum, in China, at the Lower Congo, in ]Mada- gascar, and in central Asia. Boettger's first herpetological paper is in 1869; but his catalogues of the collections in the Senckenberg Museum (1892-1898) place him clearly in the school of Boulenger. He contriljuted the account of the amphib- ians and reptiles, a volume of 826 pages, to the third edition of Brehm's Tierlehen (1892). The Modern Era of Herpetology in Europe Boettger was succeeded at Frankfurt by Robert Mertens (b. 1894), who has been an active field student in the East Indies and West Africa, as well as in the 7. By Lonnberg, Andersson, Rendahl, and Vols0e, to name only a few. 604 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES West Indies and Centra] America, with a monumental review of the lizard family Varanidae (1942) and a general account of insular reptiles as notable contribu- tions, in addition to the reports on his own expeditions. He falls sharply out of the Boulengerian school in Die Amphihien und Reptilien Europas (1928, 2nd ed. 1940), a check list drawn up on the plan of the check list for North America of Stejneger and Barbour, produced in collaboration with Lorenz Miiller. The Zoologische Sammlung des Bayerischen Staates, the repository of the Spix and Martins Brazilian collections, an independent center of herpetological studies, renewed active herpetological work with the appointment of Lorenz Miiller (b. 1868) as curator of reptiles (about 1906). Miiller was immensely stimulated by a zoological expedition to the region of the Lower Amazon in 1909; his background as a competent zoological artist, curiously enough, does not appear in his own publications. In 1932 he was joined by Walter Hellmich, who had returned from Chile with large collections and brought to herpetological studies the background of a training at the Zoological Institute of the University, thus again marking the end of the era of Boulengerian dominance in Europe. Another German center of herpetological studies was created at Magdeburg by Willy WollterstorfP (b. 1864, d. 1943), to whom lifelong deafness seems to have been a stimulus rather than a handicap. After early paleontological papers he began to devote himself more and more to the salamanders, which are so richly represented in Europe, and which lend themselves so well to observation of habits in captivity. Wollterstorif is succeeded in these interests by students and colleagues in Wolf Herre (Kiel) and Giinther Freytag (Berlin). The continuing interest in the insular lizards of the Mediterranean Islands, at first mainly a matter of nomenclatorial rivalry, has been shared by most of the herpetologists of the European continent. Even as early as the 'seventies, Theodore Eimer (b. 1843, d. 1898) called attention to the problems of environ- mental effect and of the origin of species and subspecies. Papers by Wettstein, Miiller, Mertens, and Eisentraut are written from the more modern viewpoint of an interest in speciation. The somewhat parallel insular phenomena in the West Indies and in the Gulf of California have long attracted American herpe- tologists. It seems proper to record the failure of one ambitious plan of attack on this problem in the West Indies. In conversations on West Indian herpe- tology between myself and G. K. Noble, which began in 1916 (and resulted in Dr. Noble's expedition to Hispaniola in 1922), we agreed that only direct comparison of living lizards, in good series, would be adequate to establish the degrees of differentiation from island to island; that preserved collections from different dates and scattered localities would not serve; and that only a special expedition in a suitably small vessel would answer our needs. When Gilbert C. Klingel ap- peared as volunteer aid in the Department of TIerpetology, the matter was laid before him; and the result was the perfectly planned and completely disastrous voyage of the yawl Basilisk in 1930, the story of which is recorded by Klingel in Inagua (1940). The Basilisk was fearfully storm-beaten and piled up as a total loss on the reefs of Inagua Island in the Bahamas at the very beginning of her maiden voyage. BouTenger so dominated his Continental colleagues that his influence among them persisted long after his retirement. Neither Boulenger nor his catalogues ever gained any corresponding respect in North America, which has produced SCHMIDT: HERPETOLOGY ^05 its own school of herpetology.® It is remarkable, however, that Boulenger's influ- ence should have disappeared so abruptly in London with his retirement; when his post became vacant, it was filled by a broadly educated young Cambridge graduate, H. W. Parker (1897-), who brought quite new ideas to his studies on the collection. His revision of the catalogue of the Amphibia Salientia was under- taken on a vastly more detailed and thoughtful basis, and thus has been carried through only the families Microhylidae (Boulenger's Engystomatidae) in A Monograph of the Family Microhylidae (1934), and through most of the Lepto- dactylidae. George E. Nicholls, a young British student at King's College in London, had made a noteworthy contribution to the classification of the Salientia in a little paper on "The Structure of the Vertebral Column in the Anura Phan- eroglossa and Its Importance as a Basis of Classification" ( 1916 ) . The significance of Nicholl's suggestions was more especially elaborated by the late G. K. Noble. Miss Joan Proctor (b. 1897, d. 1931), to whom Boulenger refers (I believe with affection) as "mon eleve," would perhaps have been his choice to succeed him at the British Museum. She studied with him and aided in the Division during his last four years at the ]\Iuseum. Her somewhat precarious health prevented her being taken onto the Museum staff, and a place was found for her as the curator of reptiles in the Zoological Gardens of the Zoological Society of London. Her few herpetological papers give little clue to the extraordinary competence she brought to the planning and management of the new reptile house built at the Zoo under her regime. She thus has a secure place in the history of herpetology, in the large subject of the history of the keeping of amphibians and reptiles in zoological gardens, and in her relations with the British Museum group. A most effective ''research associate" had meanwhile appeared at the British Museum in Malcolm A. Smith (b. 1875). Dr. Smith had made a large personal collection of amphibians and reptiles, and engaged actively in herpetological studies, while attached to the Court of Siam as Court Physician.^ On his retire- ment he continued these studies and greatly expanded them in the revision of the "Amphibia and Reptilia" for the Fauna of British India. The volumes for turtles and crocodiles (1931), lizards (1935), and snakes (1943) have appeared, in addi- tion to the Monograph of the Sea Snakes (1926), which effectively brings one of the smaller families of snakes up to date from Boulenger's catalogue of 1896. Emendations and additions to the Catalogue of Snakes were made by Colonel Frank AVall (b. 1868, d. 1950 of the Indian Medical Service, who had great oppor- tunities to collect and study Indian reptiles. He interested himself especially in snakes and their habits in life, and in the treatment of snake bite. His series of accounts of Indian snakes, with splendid colored plates, "A Popular Treatise on the Common Indian Snakes" (in the Journal of the Bombay Nat. Hist. Soc, 1906-1919) was unfortunately never published in book form; he is known espe- cially for his Snakes of Ceylon (1921) and for The Poisonous Terrestrial Snakes of Our British Indian Dominions (4th ed., 1928). It is something of a curiosity, more especially in a herpetological career that was essentially that of an amateur, 8. The treatment of the North American herpetological fauna is, in fact, one of the weakest features of the Catalogues. 9. See Malcolm Smith, 1946, A Physician at the Court of Siam. London: Country Life Ltd., 164 pp., illus. 606 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES that Colonel Wall should have dropped herpetologieal investigation and publica- tion completely on his retirement in 1925. The rather inflexible organization of the British Museum staff, which assigns a clerk to a Division, but does not envisage the advancement of a clerk to a curatorship, is in sharp contrast with American Museums where every office boy aspires to be director. It is thus gratifying to an American observer that J. C. Battersby (1901-), Clerk in the British Museum's Division of Reptiles, was at last placed in charge of the Division, since, when Dr. Parker was made Keeper of Zoology, the Trustees of the Museum refused otherwise to fill his vacated post in reptiles, apparently on the ground that he (Parker) could fill both posi- tions. Mr. Battersby, meanwhile, has carried on the useful role of herpetologieal bibliographer for the Zoological Record, which, after Boulenger's last contribu- tion in 1904, had been compiled by Sollas, Tate Regan, C. L. Boulenger, Joan Proctor, and Malcolm Smith. Herpetology in North America Herpetology in North America had made promising beginnings by 1850, and stood on its own feet in its exploration of the North American continent. The North American Herptology of John Edward Holbrook, published in two editions, with a multitude of emendations and separate printings of individual plates between 1836 and 1842, had established the North American region as a special field. The collections of the Academy of Natural Sciences in Philadelphia had become important, and had formed the basis for numerous herpetologieal papers in its Proceedings. In 1850 Spencer Fullerton Baird (b. 1823, d. 1887) became Assistant Secretary of the Smithsonian Institution, i.e.. Director of the United States National Museum. Baird 's interests ranged over the whole field of zoology, though certainly with herpetology as his last and most permanent love. The Catalogue of North American Reptiles, characteristically perhaps, carried no further than Part I (the snakes), was prepared jointly with the young French- man, Charles Girard (1822-1895), one of the able assistants attracted to Wash- ington by Baird. Another of these was Robert Kennicott of Chicago, whose death in Alaska in 1866, at only thirty-four, was one of the calamities to North Amer- ican zoology, and more particularly to the development of natural history in the Middle West. More important than his own writings in herpetology was Baird's indefatigable encouragement of the collecting of specimens for the rapidly grow- ing scientific collections for what was to become the United States National Mu- seum. He became the second secretary of the Smithsonian Institution (i.e., its Director) on the death of Joseph Henry in 1878 and in that capacity furthered herpetology still more by the program of publication of the new Museum, whose first Bulletin appeared in 1875, though the formal designation of the Museum as a separate entity came in 1876. The importance of Baird in the history of American science, and perhaps especially to American herpetology, can scarcely be overemphasized. Together with the encouragement of collecting and his own reports on the growing collec- tions, he furthered the careers and interests of the younger American zoologists of his day. In addition to his faithful collaborator Girard and the enthusiastic young Kennicott, there were W. H. Yarrow and finally the brilliant and inde- SCHMIDT: HERPETOLOGY 607 pendent Cope. We are fortunate to have a fine biography of Baird by his long- time associate in Washington, the malacologist W. H. Dall. Baird's untiring efforts to promote the growth of the great museum he helped to found, and his unselfish furtherance of the careers of others left him less known to succeeding generations than was the brilliant but sometimes erratic Edward Drinker Cope (b. 1840, d. 1897). Like Baird, and like most other herpe- tologists of the last century. Cope worked in many fields and is remembered quite as much for his explorations and publications in paleontology and for his studies on fishes as for his contribution to herpetology. Some of his more solid accomplishments were herpetological. They include his discovery of the pro- found difference between the true frogs and the true toads in the anatomy of the shoulder girdle and sternum, which made possible the first real advance in the classification of the whole group of tailless amphibians. Museum specimens were long jealously guarded against dissection, and their classification, it was thought, should be sufficiently accomplished by the examination of external characters. Cope's discovery, which required the laying back of the skin of the breast in order to determine the classification of a specimen in hand, ran counter to museum practice. During his European tour in 1863, when he visited the Museum of Zoology of the University of Berlin under the guidance of the still conservative Wilhelm Peters, it is said that Cope carried an open penknife in his hand and surreptitiousl}' examined the pectoral girdles of genus after genus of frogs that had previously been unknown to him. These he could then place correctly into the two great series Arcifera (for those with overlapping coracoid bones or carti- lages) and Firmisternia, with the coracoids firmly anchored to a median sternum. His early paper "Sketch of the Primary Cxroups of Batrachia salientia" (1865) sets forth this cornerstone of amphibian classification (see especially, however, G. K. Noble, below) . In addition to the continuous flow of small and large papers from Cope's pen, mainly in the Proceedings of the Academy of Natural Sciences of Philadelphia and the Proceedings of the American Philosophical Society, Cope's major herpet- ological works were The Batrachia of North America (1889) and The Crocodil- ians, Lizards, and Snakes of North America, which appeared in 1900, three years after his death. Both works incorporated large blocks of manuscript descrip- tions left by Baird. The second of these works exhibits a special direction of Cope's interest, namely the extremely varied structure of the paired copulatory organs of snakes, the hemipenes, which he figured for no less than 235 species in his Classification of the Ophidia (1895). The structure of the hemipenis, though subject to recurrent parallel or convergent evolution, and thus significant mainly at the generic level, has required the attention of herpetologists interested in the taxonomy of snakes ever since. Cope's last service to American Natural History was as editor-in-chief of the American Naturalist, 1887-1897. This gave him a ready outlet for short notes and comments, as editorials and reviews. Osborn, his biographer, aptly compares him with Lamarck; Cope was indeed a "neo-Lamarckian," believing firmly in evolution, but equally in evolution through direct influence of the environment. His mind was brilliant and polemical rather than scholarly and constructive, or, for that matter, critical. It gave off ideas and published papers like a fountain; his bibliography lists no less than 1,395 titles. Allowing for all his carelessness 608 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES JOHN VAN DENBURGH LEONHARD STEJNEGER CHARLES L. CAMP LAURENCE KLAUBER SCHMIDT: HERPETOLOGY 609 and errors, Cope was the most stimulating figure in North American zoology in the last half of the nineteenth century. Leonhard Stejneger (b. 1851, d. 1943) is the next great figure in American herpetology, contrasting as sharply with Cope as with Boulenger. He would have been mainly a contemporary of Cope's, had it not been that he came late to herpetology, at the age of thirty-eight, with a distinguished ornithological career behind him, and that Cope died at fifty-seven, whereas Stejneger lived to be ninety-one. The two herpetological careers overlapped for eight years, not without resounding clashes. Leonhard Hess Stejneger was born in Bergen, Norway. He was educated in the schools at Bergen, by private tutor, and at the University of Kristiana. He first studied medicine, in order to take the courses in zoology and botany; when he found the prospect of a medical career not to his liking, he planned to go into the family business and he entered the school of law and graduated in 1875 as cand. jur. When the business failed, he determined to make a profession of zool- ogy, instead of a hobby; and as there were few opportunities for positions in this field in Europe (let alone Norway), and on the advice of friends, he emigrated to the United States. This was in 1881; he went directly to the Smithsonian Institu- tion, and seems at once to have been given temporary employment in the National Museum by Baird. His first eight years of work were in the field of ornithology, to which he contributed a notable series of reports and the excellent volume for birds in the Riverside Natural History. This period also included his field work, financed through the United States Signal Service, on the Commander Islands, and this left an indelible stamp on his interests, as may be seen from his ambitious and sound plan for the exploration of eastern Asia (1902), his effective contri- butions to the herpetology of China and Japan, his participation in the work of the Fur Seal Commission, and his life of Steller. In 1889, on the resignation of H. C. Yarrow from the curatorship of herpetol- ogy at the National Museum, Stejneger was persuaded to take charge of this Division, and turned his attention thereafter almost exclusively to the systematics of amphibians and reptiles. He took an active part in the early field work of the United States Biological Survey in the western United States, made small collec- tions in Japan in 1896 and 1897, and took part in a collecting expedition in Puerto Rico in 1900. Thereafter he devoted himself more and more to the description of the collections flowing into the National Museum from miscellaneous sources. He was made Head Curator of Biology in 1911. The Division of Herpetology is now in charge of Dr. Doris M. Cochran (1898-), who came as Aid in 1919. Next to the IlerpetoJogy of Japan (1907), Stejneger's largest herpetological works were The Poisonous Snakes of North America (1895), The Herpetology of Porto Rico (1904), and a paper summarizing the Chinese collections in the National Museum. His smaller papers were devoted to the fauna of the United States, Mexico and Central America, the Philippines, and the West Indies, with a few from Africa and South America for good measure. His descriptions of new species are models of formal taxonomic work. When Cope produced his volume on the crocodilians, lizards, and snakes of North America, the turtles had been reserved for a separate monographic report by the comparative anatomist Georg Baur (b. 1860, d. 1898). Baur came to the United States in 1884, and by the decade of 1890 had become the leading student 610 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES of the morphology and evolution of the turtle group. He made a productive and stimulating expedition to the Galapagos Islands in 1891, and joined the staff of the University of Chicago in 1892. When Baur returned to Germany to die, Stejneger took over the project for a major work on the turtles of North America. As his time became absorbed in administrative duties, the turtle volume received only desultory attention; but he was never able to bring himself to give it up, and as result the great collection of these creatures at the United States National Museum was unavailable to other students for forty years. The monographic volume on turtles is still a desideratum. Stejneger's scholarly and critical mind was disturbed by the looseness of description of species, the failure to designate type specimens and type localities, and the indifference to orderly rules of nomenclature exhibited, in quite con- trasting ways, by both Boulenger and Cope. He introduced into descriptive herpetology the meticulous description of single specimens, which has proved to be disastrous for a usable taxonomy in the hands of some of his followers. Stej- neger never explicitly recognized the "newer theory of taxonomy as a system of group concepts based on inferences about populations from samples."^" The Boulengerian description of the species was a "paradigm" (to take over a gram- matical term) ; and is implicit in Simpson's employment of the term "hypodigm"^^ for the sum of type material available to the describer or redescriber. Malcolm Smith comments on this problem in the first of his volumes of the Fauna of British India. The lesson from Stejneger of careful designation of type specimens and of type localities, the essence of his method, was an essential advance in descriptive technique. The need for the establishment of uniform international rules of zoological nomenclature seems to have come into focus at the Fourth International Congress of Zoology, at Cambridge, England, in 1898. Stejneger attended this meeting on the occasion of his first return to Europe, which had carried him to his birthplace on museum business, and was elected a member of the first commission for nomen- clature. He became increasingly involved in nomenclatural discussion and debate, and in the succeeding meetings of the Zoological Congresses. The interest in nomenclature, and still more his treatment of its problems, seem to reflect something of the legal training of Stejneger's youth. The most con- structive herpetological result of this interest was the Check List of North Amer- ican Amphibians and Reptiles, in which Thomas Barbour joined as junior author. The five editions of this work, 1917 to 1943, witnessed a development of American herpetology and a multiplication of American herpetologists quite beyond pre- diction. Leonhard Stejneger was the last herpetologist who can be thought of as domi- nating the field for a long generation. It is characteristic that the legion of his heirs should be so numerous, so much equals, and on the whole so cooperative. The remaining history of herpetology in North America is a history of the establish- ment of active herpetological work at a whole series of nationwide centers, some- times with whole groups of active graduate students pursuing "problems." 10. George Gaylord Simpson, 1940, "Types in Modern Taxonomy," Amer. Journ. Sci., 238:417. 11. lUd. schmidt: herpetology 611 Herpetology at University Museums As a contemporary of Baird, and thus at the beginning of our century of her- petology, Louis Agassiz (b. 1807, d. 1873) appeared upon the New England scene and set in motion the greatest of university research museums, the Museum of Comparative Zoology at Harvard. Among Agassiz' varied interests, the turtles held high place; but this was equally, perhaps more, for their embryology than for their systematics. The prestige of Agassiz in America, as teacher and as exemplar of the "savant," was something America greatly needed. It had been only too easy to poke fun at the impractical and ridiculous, or even ludicrous, Rafinesque, and naturalists of sounder mind have been ridiculed by the practical ; one may recall with shame the portrait of a naturalist set forth in the last of the Leatherstocking tales, 27ie Prairie. Agassiz was no such naturalist — farmer and merchant and stagecoach driver, woman of fashion and bluestocking, college pro- fessor and schoolboy, all instantly fell under the spell of his greatness, which consisted, in fact, in his ability to convince them all of the greatness of natural history. An idea of the prestige of Agassiz at Cambridge in the eighteen-fifties is to be gained from the examination of the four volumes of his Contributions to the Natural History of the United States of America, with their superb litho- graphic illustration, and from the list of private subscribers who made possible the publication of so ambitious a work. The reader should not miss the bit of "inside dope" happily preserved by Dallas Lore Sharp in "Turtle eggs for Agas- siz" (in The Face of the Fields, 1911). Agassiz left a research museum as his greatest legacy to his adopted country, under the direction of his son Alexander, in many little known respects a greater man than his father. Louis Agassiz had himself accumulated great herpetological collections for the new Museum of Comparative Zoology — collections from the Amazon, for example — literally by the barrel. The zoologist who fell heir to these riches, Samuel Garman (b. 1843, d. 1927), after some notable contributions to the herpetology of North America, the West Indies, and the Galapagos, turned his attention more and more to studies of fishes. No full time herpetologist appeared at the "M. C. Z" until Thomas Barbour (b. 1884, d. 1946) took over the curator- ship of the division in 1910, while still a graduate student at Harvard. He became director in 1927. His interest in foreign travel, and especially in animal geog- raphy, had been whetted by diligent boyhood reading of Wallace and Bates, Belt and Hudson. After a prolonged wedding trip through the East Indies in 1906, he devoted himself more and more to the West Indies and Panama. In the Canal Zone he was perhaps more than any other person responsible for the preservation of Barro Colorado Island as a natural laboratory — and as a living exhibit of the tropical forest made accessible to the biologists of the United States (and of the world) — either for a glimpse of its magnificent plants and animals, or for pro- longed study. Barbour's large frame and booming voice, together with the pres- tige of his wealth and influence, made him a dominant figure wherever he ap- peared. It was necessary to know him more intimately to appreciate his generous and soft-hearted and often emotional side. Wealth could not save him from bitter and undeserved blows of fate. His herpetological work suffered from a readily forgivable overconfidence in his own powers. He literally worshiped Leonhard Stejneger, and joined him happily in the production of the successive editions of 612 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES the Check List. His autobiographical A Naturalist at Large (1943) gives a vivid view of "T. B.'s" immensely interesting personality. Barbour placed the curator- ship of the herpetological collection more and more in the hands of a young Eng- lishman, Arthur Loveridge (b. 1891), already with long experience in Africa. Since 1924 the collections have benefited from Loveridge's competent field work in Africa and Australia. Loveridge's published work has caught up some of the accumulations of knowledge since the time of Boulenger's Catalogues. While Curator of the Zoological Museum at the University of Michigan, Alex- ander G. Ruthven (b. 1882) had occupied himself with studies of local herpetol- ogy, and in 1906 had engaged on an active field expedition, with an essentially ecological outlook, in New Mexico, under the auspices of the American Museum. He then addressed himself to the revision of an American genus of snakes, Tham- nophis (the garter snakes), the taxonomy of which had been left in hopeless con- fusion by Cope. Perhaps mainly encouraged by Stejneger, Ruthven undertook the study of what then seemed an enormous material, drawing upon Raymond Pearl for advice as to biometric method and, by 1908, producing a measure of order in what proved to be, by example, a work of the most crucial influence in subsequent herpetological studies in America. This was his Variations and Genetic Relations of the Garter-Snakes published as Bulletin 61 of the United States National Museum. By limiting his field to a single well-defined genus, Ruthven set a pattern for further revisionary studies that lent themselves to a new mode in herpetology, the Ph.D. thesis. The University of Michigan itself, under Ruthven 's directorship and rejuvenation of its museum program, became the leading center of herpeto- logical training at the level of the university graduate school. Such university- fostered research is clearly the major herpetological phenomenon in America during the first half of the twentieth century. The succession of herpetologists in the University Museum at Ann Arbor was via Helen Thompson Gaige, long herpetological editor of the journal Copeia, to Norman Hartweg and Charles F. Walker, witli the continuing association of L. C. Stuart. In another direction the Michigan School, derived directly from Ruthven 's regime at the University Museum, leads to the scholarly Frank N. Blanchard (b. 1888, d. 1937) and to his aid and friend, Howard K. Gloyd (b. 1902) who subsequently became director of the Chicago Academy of Sciences, which tlius developed as a center of her- petological studies and publication. William H. Stickel (b. 1912) of the United States Fish and Wildlife Service affords another example of the competent train- ing of the students who came under Blanchard 's influence. During Ruthven's regime the reorganization of the University IMuseums (Paleontology, Botany, and Anthropology were combined with the Museum of Zoology) as a separate university department was realized, both in organization and in a separate new building. That the separation of the museum from the teaching departments associated with it is of vital importance is shown by the fate of departmental collections in colleges and universities the country over. That fate has been neglect, dispersal, sale, or total loss, as the heads of depart- ments changed. Revitalized museum programs in universities, or the establish- ment of new ones, in more or less conscious imitation of the museum developments at Harvard and Michigan, have been almost a sign of the times, though some universities have continued to dispose of their collections, which have frequently SCHMIDT: HERPETOLOGY 613 gone to the large public museums. Notable herpetologieal centers at universities, with teachers and graduate students in tliis field, have flourished at Cornell, Rochester, California, Florida, Illinois, Iowa, Kansas, Louisville, Texas, Tulane, Colorado, Brigham Young, Utah, and the College of Puget Sound. Among university museums maintaining expanding research collections, the high level of systematic studies at the Museum of Vertebrate Zoology at the Uni- versity of California at Berkeley requires mention. Joseph C. Grinnell, the great first director, took part in studies on the amphibians and reptiles of California and furthered the work of Charles Lewis Camp (b. 1893) on the California fauna. Grinnell and Camp now have an able herpetologieal successor at Berkeley in Robert C. Stebbins, Jr., and with Raymond C. Cowles at the University of Cali- fornia at Los Angeles, Tracy Storer at the Agricultural College at Davis, and George S. Myers and a group of active students at Stanford University, Califor- nia, has produced and is producing an active herpetologieal group, which has followed up the earlier work of van Denburgh, to be mentioned below. Herpetology in American Public Museums The larger public museums, with their dual organization as instruments of public education and institutes of research, continue to be the major centers of systematic herpetology. Such endowed museums are an especially American phenomenon, though notably represented in Europe by the Natur-Museum of the Senckenbergische Naturforschende Gesellschaft, at Frankfurt am Main. At the oldest of these in America, the museum of the Academy of Natural Sciences of Philadelphia, herpetology unfortunately failed to receive support after the death of Cope, whose herptological collections were left to the Academy. Arthur Erwin Brown (b. 1850, d. 1910), of the Zoological Society of Philadelphia, served use- fully as interim aid. Emmett Reid Dunn (b. 1894), from near-by Haverford College, himself in some respects not unlike Cope in fertility of mind, has long served the Academy as honorary curator of herpetology; but the Cope Collection needed and deserved a full-time herpetologist as curator. The decline of herpetol- ogy at the Academy came during the period of most active expansion of the field in Washington and New York. The importance of the United States National Museum to American herpetol- ogy has already been outlined. The American ^Museum of Natural History in New York City came late to an independent Division of Reptiles. Its first curator was Mary Cynthia Dickerson (b. 18G6, d. 1923), whose reputation was made by her Frog Book (1906), with its competent photographic illustration by herself. The slenderness of her subsequent herpetologieal output must be understood in the light of her creation of the first significant museum magazine, the journal now known as Natural History. Her herpetologieal importance must be weighted also for her furtherance of the careers of a succession of young naturalists — Charles Lewis Camp, Emmett Reid Dunn, Gladwyn Kingsley Noble, and myself. Noble succeeded her as Curator of Herpetology, as I believe she had planned. G. K. Noble (b. 1894, d. 1940) had been exposed equally to the influences of the Museum of Comparative Zoology and the laboratories of the Department of Zoology at Harvard and to the anatomical and phylogenetic school of William King Gregory at Columbia. He brought to the museum curatorship in New York 614 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES the plan to graft laboratory methods on taxonomic procedure, and to expand the work of his division into other aspects of natural history than the purely sys- tematic. With lavish financial support from trustees of the museum, he created a new department of ''Biology"; but he could never bring himself to give up the curatorship of herpetology. His contribution to experimental biology lay in acquaintance with and use of novel experimental animals. His contribution to taxonomy consisted in the application of Nicholl's suggestions as to the classifi- cation of the frogs, with renewed anatomical and developmental studies. Animal behavior and animal psychology led him into studies on fishes, and to the applica- tion of ideas from bird-study to herpetology, especially to courtship in lizards (1933). His most important work. The Biology of the A7nj)hihia (1931) well expresses the breadth of his interests. Noble's long succession of herpetological assistants (not to mention those in biology) began with myself and ended, at his sudden death, with Charles M. Bogert (b. 1908), with our jointly valued friend Clifford H. Pope (b. 1899) at about the middle of the series. Bogert has happily continued the tradition of a welding of experimental and anatomical techniques into a "new systematics." The Carnegie Museum in Pittsburgh built up herpetological collections, be- ginning with the Haseman expeditions to South America (primarily for fishes), and has maintained a Division of Herpetology under the curatorship of M. Gra- ham Netting (b. 1904) since 1925. At the Chicago Natural History Museum (then Field Museum) a Division of Reptiles was organized by myself in 1922. This has been under the curatorship of Clifford H. Pope (b. 1899) since 1941. In the West the public museum as research institute is represented only by the museum of the California Academy of Sciences at San Francisco. This institu- tion has had a distinguished herpetological program since the eighteen-nineties. The publishing career of Dr. John van Denburgh (b. 1872, d. 1924) extended from 1894 to 1924. He was effectively aided by Joseph R. Slevin in building up the collection, the domain of which was envisaged as the Pacific Ocean and its bordering lands. Notable in the history of the Academy was the definitive collect- ing in the zoologically classic archipelago of the Galapagos Islands. The Academy has also taken the lead in the exploration of the Lower California Peninsula (Baja California). Several of the larger museums and various university museums of the United States have engaged in the exploration of Mexico and Central America, which naturally invite the interest of herpetologists. Building upon the works of Bocourt and Giinther, our knowledge of Mexican herpetology in particular has been brought to the advanced state in which check lists of the fauna could be prepared. Check lists of the snakes (1945) , amphibians (1948) , and of the remaining reptiles (1950) by Hobart M. Smith and Edward H. Taylor summarize their own work and that of others. The Canadian fauna of amphibians and reptiles being relatively impoverished, herpetology has been little more than an appendage to the active studies, on other groups of vertebrates, that have long flourished in Canada. Herpetological col- lections have nevertheless accumulated, especially at the Royal Ontario Museum with E. B. S. Logier, at the Provincial Museum of British Columbia under G. Clifford Carl. This fauna has been supplied with a check list by R. Colin Mills (1948). schmidt: herpetology 615 Herpetology in Zoological Gardens The relations of zoological gardens with the main currents of herpetological thought depend on the personalities involved. The reptile house, at every zoo, it is said, is next only to the monkey house in popular interest. Thus a curator of reptiles is a necessity for every zoo staff, and these are usually drawn from the host of amateur snake-keepers, who, in America, replace the lizard-lovers of Europe. Thus it is natural that the herpetologist at a zoo should find himself involved in popular writing and, vice versa, the zoo job has an attraction for the snake-keeper with a flair for newspaper writing; these relations are exemplified in the career of Raymond Lee Ditmars (b. 1876, d. 1942) long curator of reptiles at the Bronx Zoo of the New York Zoological Society. For twenty or more years Ditmars' books were as books of the Bible to aspiring young herpetologists in the United States, to the dismay of those of us who saw their grave defects — that they treated herpetological knowledge as a closed book, instead of as the mere begin- ning of knowledge; that they made it seem that herpetology began with Ditmars; and that they encouraged the idea that the whole duty of a herpetologist lies in repeating a modicum of knowledge as a kind of patter, on all possible occasions. In these respects Ditmars' The Beptile Book (1907) fell far short of Miss Dicker- son's Frog Book. The appearance of more serious handbooks for the young, and especially of handbooks that suggest things to do and things to observe, now defi- nitely relegates the Ditmars era to the past. These newer books may be listed in order. For the United States, at least, it is to be hoped that they will stimulate a new period of herpetological investigation, in new and varied directions, as did the Erpetologie generaJe a hundred years before. 1933. Handbook of Frogs (3rd ed., 1949), by A. H. and A. A. Wright. 1937. Snakes Alive and Hoio They Live, by Clifford H. Pope. 1939. The Turtles of North America, by Clifford H. Pope. 1941. Field Book of Snakes, by Karl P. Schmidt and D. Dwight Davis. 1943. Handbook of Salamanders, by Sherman C. Bishop. 1946. Handbook of Lizards, by Hobart M. Smith. 1952. Handbook of Turtles, by Archie F. Carr. Ditmars' position in New York has been filled by a member of the new American school of professionally trained herpetologists, J. A. Oliver (b. 1914), lately of the American Museum and the University of Florida. As the London Zoo brought the much too sessile Boulenger into contact with living amphibians and reptiles, the great zoo at Berlin, though never with a pro- fessional herpetologist as curator, was a source of the fine photographic illustra- tion of Brehm's Tierlehen. The Cairo Zoological Gardens were long in charge of Major S. S. Flower, whose lifelong herpetological interests continued after his retirement to England. American zoos have been fortunate in their strong herpetological sections. Roger Conant carried much of the infiuence of the Michigan school from Toledo to Philadelphia, where he continued the precedent of scientific studies set by A. E. Brown. In San Diego C. B. Perkins and C. E. Shaw have made excellent use in the favorable climate of Perkins' design of a reptile house, which is quite as effective in San Diego as is the museum-type building, designed by Miss Procter, in London and Washington. An American phenomenon, the so-called "Snake Farm," has grown up in 616 A CENTURY Of PROGRESS IN THE NATURAL SCIENCES response to the great interest in snakes on the part of the general public. This is to no small degree a modern counterpart of the performances of the North Afri- can and Oriental snake-charmers. It might be passed over without mention here were it not that the Florida Reptile Institute, under the able showman E. Ross Allen, has developed via a business of herpetological supply into ambitious her- petological research. In the serpentaria of the institutes manufacturing antivenin as a remedy for snake bite, the collections of living snakes yield a by-product in the form of snake shows that correspond exactly in an inverted relation to those of the snake farm, as in Sao Paulo, Brazil, Bangkok, Siam, or Port Elizabeth, South Africa. The Amateur in ITerpetology Natural history has always been open to amateurs and self-education in this field has often preceded book knowledge. The positions in public and university museums are so few that the few actual professionals in herpetology have always welcomed the aid of volunteer students. The enthusiastic amateur needs only to follow the Huxleyan motto tenax propositi to be able to vie with professionals at their own level. It is for the amateur and beginner that the general popular books are written. Catherine C. Hopley's Snakes: Curiosities and Wonders of Serpent Life (1882), written, curiously enough, by an Englishwoman caught in South Carolina by the Civil "War, helped to set the pattern for Mary CjTithia Dickerson and Raymond Lee Ditmars. At the more serious level, it may be remembered that the only education in zoology available a century ago lay in the preparation for a medical career. Thus medical men were long the principal leaders in herpetology as in natural history in general. One may wish that the avocation of natural history studies had per- sisted as a custom among medical men, to whom studies in the field would combine recreation with the promotion of science, and to whom comparative anatomy would be a readily opened book. A late exemplar of the happy combination of a medical career with a life-long interest in herpetology was the distinguished and remarkable Howard A. Kelly (b. 1858, d. 1943), Professor of Gynecology at Johns Hopkins University. Among our colleagues of 1950 it is refugees with a Euro- pean medical training that take up functional comparative anatomy. My two American correspondents who pursue both herpetology as such and the practice of medicine are Dr. Murray L. Johnson of Tacoma, Washington, and Dr. Fred- erick A. Shannon, of Wickenburg, Arizona. The amateur who reaches the highest professional standards is likely to bring a fertilizing element of originality to his work. The most conspicuous illustration of the herpetological amateur turned professional in America is the career of Dr. Laurence M. Klauber (b. 1883). Beginning with desultory collecting of living snakes and lizards for the San Diego Zoo, he was led first to systematize his obser- vations during automobile travel at night. As night collecting proved to be vastly productive, often of species previously regarded as rare, Klauber began to build a great personal collection; as this grew, he pioneered in methods of statistical study of variation in snakes, a natural turn of interest on account of his mathe- matical training as an engineer. In the last decade of our history he was at work on a monographic account of that most distinctive of American snakes, the rattle- SCHMIDT: HERPETOLOGY 617 snake. His contribution to systematics in the fauna of the American Southwest consists in reviewing genus after genus in terms so much more exact than in any earlier work as to be beyond comparison. These studies have supplied secure foundations for further studies in any direction, which is a major function of taxonomic zoology. Regional Schools op Herpetology With the Boulengerian Catalogues available, independent schools of herpetol- ogy could grow up in South Africa, Australia, and South America. In the Union of South Africa the existence of a great number of regional museums greatly furthered the independent growth of herpetology focused on the rich fauna of the region. At the Albany Museum in Grahamstown, John Hewitt's papers begin in 1909, and two books by Walter Rose of Cape Town, Veldt and Vlei (1929) and The Reptiles and Amphibians of Southern Africa (1950), afford an introduction to this fauna at the popular level. Among numerous able students, Vivian F. FitzSimons (b. 1901), at the Transvaal Museum, took the lead with his volume on The Lizards of South Africa (1943). A Guide to the Snakes of Uganda (1938), by Captain Charles R. S. Pitman, with excellent colored plates, ingeniously financed by subscription, represents still another competent work by an amateur. In Australia an independent center of herpetology grew up at the Australian Museum in Sydney under J. R. Kinghorn. The existence of the museums of the several states in Australia has furthered publication and popularization, as in South Africa. In South America, herpetology has flourished mainly in the Argentine at Buenos Aires and in Brazil at Rio de Janeiro and Sao Paulo, with immigrant scholars from Europe, and with European and North American trained native students. j\Iiguel and Kati Fernandez, in the Argentine, have produced an excel- lent account of life histories of frogs, and Bertha Lutz, drawing upon her own and her father's notes, has taken the step from taxonomy to ecology at Rio. The work of Afranio do Amaral, long director at the Instituto Butantan, has been mainly on lizards and snakes. The extraordinary life history of Darwin's frog, in which the tadpoles are brought to maturity in the vocal sac of the male, was worked out by Karl Pflaumer in Chile between 1926 and 1930 ("Beobachtungen an Rhinoderma darwinii," Zool. Garten [1934], n.s., 7:131-134). The Brazilian group of herpetologists is especially strong at the half-century mark in 1950. Tlic Philippine fauna, after the acquisition of the islands by the United States in 1898, became tributary to the United States National Museum, and was further exploited herpetologically by the active collecting and publication of E. H. Taylor — quite in the pattern of the European colonies, but with the summary volumes published by the Philippine Bureau of Science. The independence of Chinese herpetology from European and American cen- ters was forecast before the drawing of the "Bamboo Curtain" by the work of C. C. Liu, beginning in 1930 and culminating in his large work on The Amphib- ians of West China (1950). Dr. Liu had the advantage of close relations with his American herpetological colleagues, and could build on the work of his teacher Dr. Alice M. Boring (b. 1883) and on the contributions to the herpetology of China of Clifford H. Pope. 518 ^ century of progress in the natural sciences Herpetological Societies and Journals The growth of the herpetological societies that maintain journals as outlet for publication had a most important influence on the rise of herpetology in America in the twentieth century. Copeia, now the journal of the American Society of Ichthyologists and Herpetologists, begun in 1913, was at first edited and privately published by John T. Nichols, of The American Museum of Natural History. Its function as envisaged by him was to serve as an outlet for short papers on mis- cellaneous minor observations of all sorts on cold-blooded vertebrates. The journal was taken over by the Society in 1924 under the editorship of E. E. Dunn, and was expanded and reorganized to publish longer and more important papers in 1930, under the editorship of Helen Thompson Gaige. I served as herpetological editor from 1937 to 1950, followed by Norman Hartweg. The miscellaneous note section continues the tradition of minor notes, often by beginners in the field, and thus has served as an effective training school for the writing of papers. Research is fostered by grants-in-aid from the Society's funds. Like Copeia, the journal Herpetologica was at first privately published by Major Chapman Grant, of San Diego; it was founded in 1936, and was edited by Major Grant and Walter L. Necker until 1943, subsequently by Major Grant alone. The Herpetologists' League was oganized in 1946 in order to strengthen support for Herpetologica. It is gratifying to note the birth of the British Journal of Herpetology, in 1948, as the organ of the newly organized British Herpetological Society, The infiuence of both societies and journals has plainly been to expand the numbers of herpetologists, to fire more and more amateurs with the ambition to publish their studies and observations, and to direct an increasing number of students into university training. Anatomy^ - Interest in the anatomy of amphibians and reptiles was split three ways during the century under discussion. Simple description of the anatomy of animals (and plants) has always been one of the main duties of morphologists, and this ele- mentary recording of facts continued throughout the century. The taxonomy of the higher categories is based almost entirely on morphological differences and similarities, and the pursuit of taxonomic interests added greatly to our knowl- edge of the anatomy of amphibians and reptiles. Far more important than either of these was the enormous stimulus to anatomical research that came from the publication of the Origin of Species. Amphibians and reptiles occupy a strategic position between the fishes and the mammals, and were closely studied in the intensive search for the phylogeny of vertebrate structures. In 1850 the field of vertebrate anatomy was still dominated by the methods and ideas of Cuvier in France, Meckel and Johannes Miiller in Germany, and Owen in England. The works of these four and their contemporaries, aside from their philosophical content, laid the foundation for modern descriptions of the anatomy of vertebrates. Straight description of structure, perhaps because it usually does not attempt to evaluate data and therefore demands little back- ground, is available doctor's thesis material. Many of the hundreds of anatomical 12. Contributed by my colleague, D. Dwight Davis, Curator of Anatomy, Chicago Natural History Museum. SCHMIDT: HERPETOLOGY 619 papers published during the past century are by obscure persons who are never heard of again, or by men who later became noted in nonmorphological fields. A few, however, stand out because of the number or importance of their studies. Owen himself carried over into the post-1850 period. The first volume of his Anatomy of Vertebrates, which covers amphibians and reptiles, appeared in 1866 when Owen was sixty-two years old. St. George Mivart (b. 1827, d. 1900), an isolated half-mystical figure who for many years was Lecturer in Comparative Anatomy at St. Mary's Hospital in London, contributed several careful descrip- tions of the skeleton and muscles of amphibians and lizards before he turned his attention exclusively to mammals. William Kitchen Parker (b. 1823, d. 1890) was a British physician with a passionate love of nature that expressed itself in a series of meticulous monographs, illustrated with plates drawn by himself, on the structure and development of the skull and pectoral girdle of various amphib- ians and reptiles. Parker was greatly handicapped by the fact that he could not read German. That remarkable Swede, Gustaf Retzius (b. 1842, d. 1919), could hardly have failed to contribute to our knowledge of the morphology of amphibians and rep- tiles. Retzius was the son of the distinguished anatomist and anthropologist, Anders Adolf Retzius, who in turn was the son of a distinguished natural scien- tist. Retzius was a friend of the great German anatomist, Johannes Miiller. His work was almost wholly descriptive, painstakingly detailed, and illustrated largely by himself. The tremendous Das Gehororgan der Wirhelthiere (2 vols., 1881- 1884) contains meticulous descriptions of the auditory apparatus of many am- phibians and reptiles. Later, after he had turned his attention to the structure of sex cells, he described the spermatozoa of many amphibians and reptiles. The outstanding descriptive work of this era is Die Anatomie des Frosches, which was addressed to physiologists rather than anatomists. The first edition of this famous work, by Alexander Ecker (b. 1816, d. 1887) and Robert "Wiedersheim (b. 1848, d. 1923) , appeared in three parts between 1864 and 1882. Both Ecker and Wiedersheim were at the University of Freiburg, where Ecker was Professor of Human and Comparative Anatomy and Wiedersheim Extraordinary Professor. A second edition, completely rewritten by Ernst Gaupp (b. 1865, d. 1916), also of Freiburg, appeared in three parts between 1896 and 1904. An English edition of the first German edition, translated by George Haslam, was published in London in 1889. Other frog anatomies during this era were by Mivart (1874) and A. M. Marshall (1882) in England, and by S. J. Holmes (1916) in America. It is ex- traordinary that a comparable work on a salamander did not appear until 1934, when TJie Anatomy of the Salamander [Salamander maculosa], by Eric T. B. Francis, was published in England. Still more remarkable is the fact that no modern descriptive anatomy of any reptile has ever appeared. The most ambitious compendium of accumulated data on the morphology of amphibians and reptiles appeared in Bronn's Klassen und Ordnungen des Thier- reichs. The herpetological volumes, running to more than 2,800 pages and 223 lithographed plates, were published between 1873 and 1890. They were compiled by Christian Karl Hoffmann (b. 1841, d. 1903) of the University of Leiden. Although now sadly out of date, Hoffmann's is still the only general compilation of anatomical data for amphibians and reptiles. The classical treatise on the embryology of an amphibian is Goette's folio 620 A CENTURY Of PROGRESS IN THE NATURAL SCIENCES Die EnUvickhmgsgeschichte der Unke (Bombinator igneus) als Grundlage einer vergleichende7i Moi'phologie der Wirbelthiere (1875). Alexander Goette (b. 1840, d. 1922) was greatly influenced by the embryologist von Baer, and was himself a teacher of Wilhelm Roux. His Bombinator monograph was the basis for his purely mechanistic theory of evolution, which undoubtedly influenced Eoux's later mechanistic concept of morphogenesis. It is also the prototype of all later descrip- tive work on frog embryology. The second half of the nineteenth century was the Golden Age of the morpho- logical sciences. Knowledge of the structure and development of amphibians and reptiles, along with the other vertebrates, was enormously extended and deepened during this period. Carl Gegenbaur (b. 1826, d. 1903), more than any other man, is identified with this flowering of morphological interest. Darwin's evolutionary ideas were becoming current at the very beginning of Gegenbaur's career, and he grasped their significance at once, realizing that the phylogeny of vertebrate structure provided comparative anatomy with the conceptual framework that had previously been lacking. Our knowledge and understanding of the structure of amphibians and reptiles was enormously increased as a by-product of the research resulting from this reorientation. Gegenbaur himself contributed directly in a number of publications, but his indirect influence on herpetology was far more important. Among his assistants during his long career at Jena (1855-1872) and Heidelberg (1872-1900), Max Fiirbringer, Friedrich Maurer, Ernst Goppert, and Georg Ruge added greatly to the fund of knowledge, especially of the musculature and its innervation. His pupils carried Gegenbaur's ideas beyond Jena and Heidelberg, and even beyond the borders of Germany. Although the Gegenbaur tradition was never strong in England or America, his pupils Hans Gadow (b. 1855, d. 1927) in England and H. H. Wilder (b. 1864, d. 1928) and W. B. Scott (b. 1858, d. 1947) in America were active and influential in the English-speaking world. Schools of associated workers, often with special orientations and traditions that ran through several generations, were characteristic of central Europe. These begin with one vigorous personality, who infects and often dominates others. The Gegenbaur school, with its unflagging pursuit of the phylogeny of structures via interpretative homologies, has already been mentioned. The output of this school ran heavily to myology, a subject in which Gegenbauer himself was little inter- ested. The myological orientation is probably attributable to Max Fiirbringer (b. 1846, d. 1920). The Freiburg school, beginning with Ecker and Die Anatomie des FroscJies, and continuing through Gaupp and Wiedersheim, centered its attention largely on amphibians. In Vienna the towering figure of Joseph Hyrtl (b. 1811, d. 1894) began a dynasty that lasted through three generations, until it was destroyed by the Nazis in the years before World War II. Hyrtl's interest in the vascular sys- tem is strongly reflected in the work of Emil Zuckerkandl (b. 1849, d. 1910), Julius Tandler (b. 1869, d. 1936), and Anton Hafferl, and in the painstaking solu- tion of problems arising in the medical dissecting room, which repeatedly inspired extensive comparative researches based on the museum collections, and is evident in most of the Vienna studies of this era. Most of this work, which has a charac- teristic stamp, appeared in the Denkschriften and Sitzungsberichte of the Vienna Academy. SCHMIDT: HERPETOLOGY 621 Outside Europe the outstanding example of a special anatomical "school" is probably the extensive work in South Africa, by various authors, on the cranial anatomy of amphibians studied by means of serial sections. This work, which falls in the second quarter of the twentieth century, is traceable to C. G. S. de Villiers, who was a student of Arnold Lang in Ziirich. The third great drive of morphological research is to provide a basis for taxonomy. This, of course, involves studying many representatives of a group — often every available genus. Two quite different goals are involved. One is to distinguish the groupings into which species, genera, or families may be parti- tioned; this is essentially analytical. The other is to determine the interrelation- ships among these groupings; this is essentially synthetic. Although he was not an anatomist, G. A. Boulenger is chiefly responsible for the breakdown into families among the Rcptilia that is in use today. Boulenger, in turn, drew heavily on Friedrich Hermann Stannius (b. 1808, d. 1883), a Ger- man comparative anatomist who, after studying under Johannes Miiller, was professor at Rostock. The second edition of Stannius' Ilandhucli der Anatomie der Wirhelthiere (1854), which is set up in a taxonomic rather than an organ-sys- tem framework as the first (1846) edition w^as, is repeatedly cited by Boulenger. Cope is Boulenger's counterpart for the Amphibia, and the modern arrange- ment of families of salamanders and frogs is essentially that of Cope, sharpened and refined by a host of later workers. H. H. Wilder made the important dis- covery of lunglessness in certain salamanders in 1894. G. E. Nicholls, who was Professor of Biology at Agra College, Agra, India, discovered the importance of the vertebral column in classifying Salientia (1916). And G. K. Noble drew the soft anatomy, especially the thigh musculature, into a general review of the clas- sification of these animals (1923). Noble's work is further important for its emphasis on interrelationships rather than mere partitioning. Edoardo Zavattari, of the Zoological Museum of the University of Turin, pub- lished in 1910-1911 a 122-page monograph on the hyoid muscles of lizards, de- scribing and illustrating the patterns in a wide selection of species. This, plus earlier analytical work on the skeleton by Cope and others, and on the body and limb muscles by Fiibringer, Gadow, and Maurer, formed the basis for a general review of the classification and interrelationships of the lizards by C. L. Camp (1923). The foundation of the modern classification of turtles was laid by Boulenger. This was refined chiefiy by the voluminous work of Georg Baur and Friedrich Siebenrock, both of whom were active but not very imaginative anatomists. Bou- lenger was also responsible for the framework of the modern classification of snakes. Boulenger's classification has been improved and corrected by many later workers. A brief review of the comparative anatomy of snakes and its implica- tions was published as recently as 1951 by Bellairs and Underwood in Biological Reviews. The outstanding student of the eye of reptiles was Gordon Lynn AValls (b. 1905), who built on the earlier work of the German, Victor Franz. Walls, formerly at Wayne University at Detroit and now at the University of California, emphasized the profound differences between the eyes of snakes and lizards, and made this the basis for his theory of the origin of snakes from noctural lizards. He described, among other things, the existence of physiologically yellow lenses that 622 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES function as color filters for increasing visual acuity. All of Walls's work reflects a lively interest in functional mechanisms rather than static structure. His major work is The Vertebrate Eye and Its Adaptive Radiation, published by the Cran- brook Institute of Science in 1942. Comparative functional anatomy, in which the description of adaptive mech- anisms drives the student from the dissecting table to the living animal in the field and from the field or zoo to the dissecting table, is a relatively new direction of interest in verteberate anatomy. The phylogeny of adaptation may be pursued (as by Walls) and an understanding of the structures involved is often to be gained by the comparison of analogous, as distinguished from homologous, mechanisms. An outstanding representative of this fertile movement in herpetology was Walter Mosauer, a student of Franz Werner's in Vienna, who had made a notable contri- bution to the anatomy of snakes and to the understanding of their locomotor musculature before his untimely death in 1937. Mosauer had become a citizen of the United States and had taken his doctor's degree at the University of Michigan. The Study of Snake Venom The study of snake venoms forms a large chapter of herpetology. The scien- tific study of venoms and of the treatment of snakebite falls almost entirely within the period 1850-1950. An important preliminary study by Dr. S. AVeir Mitchell (b. 1829, d. 1914), in 1861, set the investigation of venoms and of the medical treatment of the bites of poisonous snakes on a critical and experimental basis. Sir Joseph Fayrer's TkanatopJiidia of India (1872) was supplemented by a series of papers on the physiological effects of the venoms of Indian snakes by Fayrer and Brunton (1872-1875), and further work was reported by A. J. Wall in Indian Snake Poisons; Their Nature and Effects (1883). The whole subject is then summarized by Mitchell and Reichert in Researches upon the Venoms of Poisonous Serpents (1886), in the Smithsonian Contributions to Knowledge. A burst of interest in the treatment of snakebite came with the discovery that antivenins are produced in tlie blood of animals inoculated with small successive doses of venom, and that the degree of immunity can be built up by successively increasing the dosage of venom. The pioneer students were H. Sewall, working with rattlesnake venom and pigeons (1887), and Maurice Kaufmann, using the venom of the European viper and the guinea pig (1889). This discovery led directly to experiments by Marie Phisalix and G. Bertrand at the Paris Museum and A. Calmette at the Pasteur Institute at Lille on the use of the blood serum of immunized animals as an antidote in snake poisoning. This set the stage for the development of institutes for the production and distribution of antivenins for general use. Pasteur Institutes were established at Calcutta and Bangkok. The Instituto Butantan at Sao Paulo, Brazil, was set up as much for research as for antivenin production. The Mulford Drug Company's antivenin division in the United States (with its successors) grew out of the interest aroused by the Antivenin Institute of America, which published a Bulletin (1927-1932). In South Africa, centers of antivenin production were developed, as at Port Eliza- beth. In Australia critical studies of snake venoms have been in progress under the direction of H. C. Kellaway since 1929. The enthusiastic interest aroused by SCHMIDT: HERPETOLOGY 623 the all-but-miraculoiis recoveries from serious cases of snake-poisoning in human beings after the injection of antivenin, together with the sound basis of fact as to immunization in general, led to great public interest and government support for such institutes. The first major difficulty to develop in the treatment of snakebite with anti- venin lies in the radical difference between the neurotoxic venoms of the cobra and its relatives and the haemotoxic venoms of the vipers and pit-vipers. This is especially complicated by the fact that the widespread South American rattle- snake, alone among the pit-vipers, has a powerfully neurotoxic venom. Further- more, it soon developed that the antivenins were in general strictly specific. The fact that the venoms of the many different species of poisonous snakes are sharply peculiar to the species, and that the antivenin prepared from inoculation with venom of the banded rattlesnake, for example, would not serve as an antidote for the bite of the copperhead, led to the production of "polyvalent antivenins." The specificity of venoms may be sharply marked even within otherwise barely dis- tinguishable races of a single species, and thus adds an example of the biochemical nature of species differentiation. The antivenin institutes, in retrospect, appear to have acquired a "vested interest" in snake bite, and their statistics are in urgent need of critical review. In 1927, Dr. Dudley Jackson, of San Antonio, Texas, found that in rattlesnake bite, incisions and suction on the swollen limb would lead to a high percentage of cures without antivenin. Afranio do Amaral, long Director of the Instituto Bu- tantan, was led to propose progressively greater dosages of antivenin, finally to the amount of 225 cc. This, on the face of it, introduces new dangers and new problems. At midcentury the subject is thus in need of a renewed objective and critical study. Experimental Physiology and Embryology The broad fields of experimental investigation into physiologic and develop- mental processes have had so great a growth in the 1850-1950 century, and their focus has been so much on the process and on the principles involved and so little on the particular experimental animal, that the history of the herpetological aspects of these sciences and their bearing on the growth of herpetology as a whole need not be elaborated here. A late bibliography of experimental embry- ology is available in Eugh's work with this title (1948). Salamanders, with their capacity for complete regeneration of limbs, have been especially favorable ma- terial for studies in regeneration (T. H. IMorgan, Regeneration, 1901; E. Kor- schelt, Regeneration and Transplantation, 1907). For a conspectus of the litera- ture on general physiology as applied to amphibians and reptiles reference may be made to Heilbrunn's Outline of General Physiology (1943) and to Comparative Animal Physiology, edited by C. Ladd Prosser (1950). The physiology of the whole animal, which relates it to its environment, is a part of ecology. Ecology and Herpetology Ecology, as natural history made critical and exact, stands in direct relation to modern herpetology, and requires thoughtful assessment of its origins and present status in this relation. Taxonomic herpetology in the Boulengerian Era 624 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES remained consciously aloof from consideration of habitats, and geographic dis- tribution was cited as if it were a matter of occurrence in space independent even of altitude. The characteristics that distinguish species were referred to as "use- ful" if they were useful to the taxonomist in his discrimination of systematic groups. The realization that species and subspecies had to be revalued and redescribed in terms of the general environments and special habitat niches in which they occur came first from the side of popular natural history (e.g., Brehm's Tierlehen) . Laboratory studies of the reactions and tolerances of animals afford another of the roots of animal ecology. A pioneer paper in tlie United States was based by Alexander G. Ruthven on field studies in the American Southwest in 1906 (Ruth- ven, 1907) , in which he was obviously influenced by C. C. Adams. Since that date there has been increasing interest in the observation of the biotic and physical environments in which amphibians and reptiles live, how they meet the adverse factors in their surroundings, and, in general, how they "behave" in relation to them. The importance of environmental observation to a definitive taxonomy is especially illustrated by the work of Henry S. Fitch on the garter snakes of the Pacific region (1940). Ecological observation, of course, stands on its own feet independent of its significance to taxonomy, and becomes increasingly inde- pendent as the taxonomy becomes mature, and thus a sound foundation for ecology. Ecology involves a vast variety of subsciences from physiography, mete- orology, and chemistry to the complex of biotic relations, and more particularly for herpetology, the relation of animal life to its plant matrix. Finally, since animal behavior rests on the interaction of internal physiology and stimulus from the environment, the ecology of animals must particularly include their behavior, the study of which tends to be distinguished as the separate science of animal behavior. Physiological investigation in herpetological ecology is to be discerned in the continuing studies of Raymond B. Cowles (b. 1896) and of his student and colleague, C. M. Bogert, on the temperatures of amphibians and reptiles in rela- tion to the temperature range of their environment. The sharpness of limitation to specific habitat niches reflects the long evolution of the reptile group; it is illus- trated by the rock-crevice habitat of such lizards as Sauromalus in the American Southwest, and especially by Xantusia henshawi and arizonae, which live under the loose exfoliating rockflakes of rounded granite boulders. Courtship behavior, with the frequent correlation of the spacing of individual animals of breeding groups into territories, is an important fleld of study pioneered in herpetology by G. K. Noble (Noble and Bradley, 1933). The sub.ject of "Home Ranges and Wan- derings of Snakes" {Copeia, 1947, pp. 127-136) is summarized by William F. Stickel and James B. Cope. That the populations of amphibians and reptiles are often vast has long been known from their breeding aggregations. Actual meas- urements of population density are extraordinarily few. Pioneering studies in this direction rest on the techniques of marking individuals by tagging, toe- clipping, scale-clipping, or tattooing, pioneered by F. N. Blanchard in 1933. Cagle's paper in 1950, "The Life History of the Slider Turtle, Pseudemys scripta troostii (Holbrook)" in Ecological Monographs (20:31-54, 18 figs.) summarizes ten years of work in this field. The study of distribution depends directly on examination of the present environment and on speculations regarding the past changes in environment, i.e., on ecology and paleoecology. Studies on defensive SCHMIDT: HERPETOLOGY 625 and warning behavior involve the assessment of the function of venoms, especially in relation to mimicry and coloration. Sense perception and orientation have been studied in relation to food capture and to such phenomena as the movement of hatching seaturtles to water. The problem of isolating mechanisms between species (voice in frogs, for example) involves restudy of the so-called taxonomic characters in order to find out what they mean. Especially significant studies on the physiological isolation of the .species of frogs and of populations of a single species have been made by John A. Moore (b. 1927) of Barnard College. An ecological framew^ork for studies of animal distribution was outlined by Richard Hesse in 1924 {TiergeograpJiie auf oekologischer Grundlage, Amer. ed. 1951). A framework by means of which past and future studies on the ecology of reptiles and amphibians can be brought into relation and correlation with other studies is provided by C. W. Allee, et al., Princi'ples of Animal Ecology (1949). At the midcentury the study of amphibians and reptiles may be seen to be in need of a world synthesis, perhaps a more elaborate one than that of the cata- logues of the British Museum two generations earlier, but still essentially a sys- tematic review. A review of the existing systematics of these groups should serve as a springboard from which the new systematics can be explored and applied, involving the reassessment of the classification from class to subspecies and population in the light of the advances in biology as a whole. The review envis- aged would then be the basis also of a neiv natural history, in which studies of life histories and habits and behavior are brought into relation with comparative functional anatomy. The new systematics and the new anatomy are essential to the interpretation of the still growing body of knowledge of the extinct forms of both amphibians and reptiles. We thus envisage a major contribution from her- petology to an understanding of the evolution of the animal kingdom, with its vast perspective in time and its broad ramifications in the present, BIBLIOGRAPHY Ar.Assiz, Elizabeth Gary 1886. Louis Agassiz, His Life and Gorrespondence. 2 vols., illus. Boston: Houghton, Mifflin Go. Anonymous 1870. Dumeril (Andre-Marie Gonstant). Grand Larousse, 6:1379. 1874. Peters (Guillaume-Gharles Hartwig). Grand Larousse, 12:703-704. 1931. Miss Joan Proctor. The London Times, Sept. 21, 1931. 1941. G. K. Noble. Gopeia, vol. for 1940, pp. 274-275, portr. 1951. Sherman C. Bishop, 1887-19.^1. Proc. Rochester Acad. Sci., Vol. 6 [portr. and text inserted without pagination before p. 327]. Eaubour, Thomas 1919. Boulenger, the man and his work. Nat. Hist., 19:566-567, portr. 1943. Naturalist at Large, xii + 314 pp., illus. Boston: Little, Brown and Go. Bakdeleben, Karl 1894. Joseph Hyrtl. Anat. Anz., 9:773-776. Bertin, Leon 1939. Gatalogue des types de poissons du Museum National d'Histoire Naturelle. Ire pte. Gyclostomes et Selaciens. Bull. Mus. Natl. Hist., Nat, s6t. 2, 11:51-98 [Historique des collections ichthyologiques du Museum, pp. 51-63]. 626 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES B[lanford], W. T. 1900. John Anderson, M.D., LL.D., F.R.S. . . . , Nature, 62:529-531. Bluntschli, Hans 1922. Max Fiirbringer. Anat. Anz., 55:244-255, portr. BOULENGER, G. A. 1906. Reptiles and Batrachians, m The History of the collections ... of the British Museum, vol. 2, pp. 517-531. London. British Museum. 1921. Liste des publications ichthyologiques et herpetologiques (1877-1920). Ann. Soc. Roy. Zool. Malac. Belgique, 52:11-88. Cattell, Jaques, Ed. 1949. American Men of Science a Biographical Dictionary. 8th ed. [iv] + 2836 pp. Lancaster: The Science Press. Dall, William Healey 1915. Spencer Fullerton Baird, a Biography including Selections from His Corre- spondence with Audubon, Agassiz, Dana, and Others, xvi + 462 pp., illus. Philadelphia: J. B. Lippincott Co. DUMEEIL, A. M. C. 1854. Notice sur G. Bibron. In Erpet. Gen., Vol. 7 [inserted 4 pp. notice after preface, unpaged]. Fischer, Albert Kenrick 1943. Leonhard Stejneger. Copeia, vol. for 1943, pp. 137-141. FrEYTAG, GtJNTHER E. 1948. Willy Wolterstorff. Ein Forscherleben fiir das Museum fiir Naturkunde und Vorgeschichte zu Magdeburg. Abh. Ber. Mus. Naturk. Magdeburg, 8:7-18, 1 pi. Gloyd, Howard K. 1940. Frank Nelson Blanchard, Scholar and Teacher. Herpetologica, 1:197-208, illus. GoDMAN, Frederick Ducane 1915. Biologia Centrali-Americana Zoology, Botany and Archaeology. Introductory Volume, VIII + 149 pp., 2 pis., 8 maps. Privately published. Gregory, William K. 1942. Gladwyn Kingsley Noble (1894-1940). Yearbook Amer. Philos. Soc, vol. for 1941, pp. 393-397. GUNTHER, A. C. L. G. 1886. Reptiles, History and Literature. Encyclopedia Britannica, ed. 9, 20:432-441. GuNTHER, A. C. L. G., and A. S. Woodward 1911. History of Herpetology. Encyclopedia Britannica, ed. 11, 23:136-141. GtJNTHER, R. T. 1930. Bibliography of the works of Dr. Albert Gunther. Ann. Mag. Nat. Hist., ser. 10, 6:233-286. Haas, [Fritz] 1910. Professor Dr. 0. Boettgerf. Ent. Blatt., 6:267-268, portr. Hay, 0. P. 1898. George Baur. Science, n.s., 8:68-71. Hertwig, Oscar 1905. Ann. Rep. Smithson. Inst, for 1904, pp. 787-791, portr. SCHMIDT: HERPETOLOGY 627 MosAUEE, Walter 1940. Franz Werner, 1867-1939. Herpetologica, 1:178-183, portr. NORDENSKIOLD, ERIK 1942. The History of Biology, xii + 629 [xv] pp. New York: Tudor Publishing Co. N[orman], J. R. 1938. Dr. G. A. Boulenger, F.R.S., Nature, 141:16-17. Osborn, Hexry Fairfield 1931. Cope: Master Naturalist. The Life and Letters of Edward Cope, with a Bibli- ography of His Writings Classified by Subject. A Study of the Pioneer and Foundation Periods of Vertebrate Paleontology in America, xvi + 749 pp., illus. Princeton: Princeton Univ. Press. PlETSCHMANN, V. 1919. Franz Steindachner. Ann. Naturhist. Mus. Wien, 33:47-48. RouLE, Louis 1917. Notice necrologique sur M. le Dr. Frangoic Mocquard. Bull. Mus. Natl. Hist. Nat., 23:315-317. RuTHVEN, Alexander G. 1931. A Naturalist in a University Museum. 143 pp., illus. Ann Arbor: Privately printed. 1936. Helen Thompson Gaige. Herpetologica, 1:1-2, portr. Schmidt, Karl P. 1936. Robert Kennicott, Founder of Museums. Program of Activities, Chicago Acad. Sci., 7:3-8. 1937. Frank Nelson Blanchard, 1888-1937. Copeia, vol. for 1937, pp. 149-150. Seab.ia, Antoxio Maria d'Albuquerque 1891. Esboco biographico do excellentissimo senhor Dr. Jacques Wladimar de Bedriaga. 8 pp. Coimbra; Imprensa da Universidade. Slye, Maud, John Oliver La Gorge, G. Kingsley Noble, and Barrington Moore 1923. Mary Cynthia Dickerson, 1866-1923. Nat. Hist., 23:507-519, portr., illus. Smith, Malcolm 1938. George Albert Boulenger, 1858-1937. Copeia, vol. for 1938, pp. 1-3, illus. 1947. Stanley Smyth Flower, 1871-1946. Copeia, vol. for 1946, pp. 185-187, portr. 1951. Frank Wall, 1868-1950. Copeia, vol. for 1951, pp. 113-114, portr. Waldeyer-Hartz, W. von 1920. Gustaf Retzius. Anat. Anz., 52:261-268, portr. Wetmore, Alexander 1945. Biographical Memoir of Leonhard Hess Stejneger (1851-1943). Natl. Acad. Sci. Biogr. Mem., Vol. 24, 4th memoir, pp. 145-195, portr. VVettstein, Otto von, Fr. Maiol, and Joseph Eiselt 1941. Franz Werner als Mensch und Forscher. Ann. Naturh. Mus. Wien, 51:8-53, portr, Wheeler, William Morton 1899. George Baur's Life and Writings. American Naturalist, 33:15-30, portr. Wood, N. L. 1944. Raymond L. Ditmars; His Exciting Career with Reptiles, Animals, and Insects. X + 272 pp., illus. New York: Julian Messner, Inc. ORNITHOLOGY Bij CHAELES G. SIBLEY Cornell University The first half of our Century of Progress was, for ornithology, concerned almost entirely with systematics. Collections were growing, new species were still being described with frequency, and the description of the unknown multi- tude of subspecies had barely begun. Most attempts to interpret the sigificance of behavior met with failure because the necessary premises had not yet been developed. The study of natural history was peculiarly typical of northern Europe. England and Germany produced a majority of the naturalists of the time. The expansion of the colonial empires of the European nations resulted in extensive travel and the establishment of many private fortunes. With the tradition estab- lished and the means available it was a logical consequence that the study of birds should prosper. Knowledge of North American birds prior to 1850, was largely due to the work of Alexander "Wilson, Charles Lucien Bonaparte, William Swainson, John James Audubon, and Thomas Nuttall. Others there were, but these five pro- duced the most extensive publications and illustrations. With Audubon's death in 1851, the pioneer era in American ornithology came to a close. By 1853 ornithology was past its infancy. Few indeed were the major areas of the earth from which collections had not found their way to Europe or America. In Germany Herman Schlegel had recently (1844a, 1844b) begun to employ trinomials to designate geographic races and in a country house in Eng- land Charles Darwin was quietly working on a book (1859) which was to ini- tiate great controversies and provide the stimulus for intensified research in all fields of biology for the next century. There is a curious parallel between the histories of two German clerics of the 1860's. Both Gregor Mendel and Bernard Altum were ahead of their time. The importance of Mendel's now famous work (1865) went unrecognized for over thirty years, while Altum 's concept of territory (1868) was not "discovered" until after H. Eliot Howard (1920) had independently arrived at similar conclusions. The last part of the nineteenth century was marked by numerous local fau- nal treatises, especially in Europe, and by the issuance of elaborate monographs on various groups of birds. At the halfway point in our century, Robert Ridg- way (1901, p. 1) epitomized the prevailing viewpoint of the time when he wrote: There are two essentially different kinds of ornithology: systematic or scientific, and popular. The former deals with the structure and classification of birds, their synonymies and technical descriptions. The latter treats of their habits, songs, nesting, and other facts pertaining to their life-histories. . . . Popular ornithology is the more entertaining, with its savor of the wildwood, green fields, the riverside and seashore, bird songs, and the [629] 630 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES many fascinating things connected with out-of-door Nature. But systematic ornithology, being a component part of biology — the science of life — is the more instructive and there- fore more important. The understanding of the true significance of what Ridgway called "popu- lar" ornithology had to await the development of a firm foundation of physio- logical, psychological, and ecological research. These in turn depended heavily (the debt is all too seldom acknowledged) upon a foundation of systematics. It was inevitable that "scientific" ornithology and "systematics" should seem sy- nonymous to Ridgway. Even as Ridgway wrote, the revolution was starting. Mortensen was band- ing birds in Denmark, Selous in England had started his energetic advocacy of the study of the living bird, and Chapman in the United States was urging the partial substitution of the binocular for the shotgun. Within the first decade of the present century banding began to solve prob- lems concerning migration. Heinroth showed how behavior could be a clue to phylogentic relationship, and Howard helped to reveal the self-deceiving pitfall of anthropomorphism. The barrier between the "scientific" and the "popular," which seemed so clear to Ridgway, was beginning to disappear. If any one year may be selected as a "turning point" it would fall close to 1920. Until then it was possil^le to function as an adequate ornithologist if one was versed in the systematics, distribution, and life histories of birds. The in- crease of interest in psychology and physiology at first concerned only a few. Most scientific zoologists saw only the shadow of the "bird lover" in ornithology. (Some still do!) The feeling that the study of birds was unimportant to the serious scientist prevailed. Since 1920 there have been great changes. Birds have come to be recognized as providing excellent material for the study of ani- mal behavior and evolution. Many of the phases of ornithology which Ridgway had dismissed as "popular" are now among the most abstruse subjects of zoology. Indeed, these items compose an important group of factors contributing evidence of relationships. In 1953 it would be impossible to justify a division into "two essentially different kinds of ornithology." The effective avian systematist of today must be more tlian a mere cataloguer with an eye for variation. Behavior, ecological relationships, physiology, genetics, and even parasites are utilized as clues to phylogeny. The recognition of evolution as the central theme of all biology has obliter- ated the sharply drawn boundaries between disciplines even as the same realiza- tion has shown the reality of the blurred lines between our arbitrary taxonomic categories. Systematics and Evolution Two factors were largely responsible for the taxonomic viewpoint of the mid-nineteenth century : the belief in the immutability of the species promoted a strictly morphological species concept, and collections were mostly too limited to reveal the full breadth of individual and geograiihic variation. The change in the point of view toward the first of these factors was to depend upon the ac- ceptance of the evolutionary doctrine and was destined to be a matter for debate during most of the next century. As for the second factor, collections were al- ready growing rapidly. SIBLEY: ORNITHOLOGY 631 Taxonomic practices during the early nineteenth century were in accord with the concepts of the time. If a newly acquired specimen differed from the "type" of the most closely related known species it became the type of a "new species." Some authors described each variant as a new species, regardless of the degree or cause of the differences. As collections grew it became apparent that not all the "species" which were being described were of equal rank. The first attempt to reflect differences in the rank of forms below the species level was made by Carl Friedrich Bruch who proposed (1828) that "variations" be designated by a third name added to the Linnean binomial. Fourteen years later, on Septem- ber 23, 1842, at the twentieth annual meeting of the Society of German Natural- ists and Physicians, Bruch (1843) again expounded his ideas before the zoologi- cal section of the society. He used the term "subspecies" and again cited trinominal combinations. At this meeting Hermann Schlegel (b. 1804, d. 1884) was the presiding officer. Whereas Bruch had used ternary nomenclature to designate any relatively slight degree of departure from the "typical," Schlegel soon began to apply it only to geographic variants. His first use of trinominals was in 1844 in the "Aves" section of Siebold's Fauna Japonica. Schlegel, al- though the junior author (with C. J. Temminck), was responsible for the nomen- clature and used such combinations as Pandion haliaetus orientalis (p. 13), Otus scops japonicus (p. 27), and Podiceps ruhricoUis major (p. 122). Schlegel also employed the trinominal in his critical review of European ornithology in 1844. The use of trinominals found no protagonists in Europe, and it was in the United States that it first gained general acceptance among ornithologists. Spen- cer Fullerton Baird (b. 1823, d. 1887) had begun the detailed ornithological ex- ploration of North America in 1850 when he became the assistant secretary of the Smithsonian Institution. The survey trips for the Pacific Railroad brought in large numbers of specimens and by 1858 Baird was able to recognize numerous examples of geographic variation in his series. The evidence for general rules of geographic variation was also found by Baird. He noted Bergmann's Rule (body size increases toward the north and decreases toward the south) and in 1859 pointed out that the same change takes place in accordance with changes in altitude in the same latitude. Baird also noted that in the southern parts of its range a species tends to show relative increase in bill size and that Pacific Coast specimens of many species were darker than those from inland localities. He was well aware of the tendency for the characters of adjacent differentiated populations to merge (i.e., intergrade) where the margins of their ranges ad- joined. In 1858, with the cooperation of John Cassin and George N. Lawrence, Baird published the famous ninth volume of reports on the Pacific Railroad sur- vey. Under Baird's skillful direction this volume became far more than a mere "report." It was in fact the most important treatise on the systematics and nomenclature of North American birds up to that time and remained so for many years. It was Baird's protege, Robert Ridgway (b. 1850, d. 1929), who next applied himself to the problem of the boundary between species and subspecies. AVhen but seventeen years old Ridgway was appointed zoologist to the United States Geological Survey of the 40th Parallel. The expedition went to Panama by ship, crossed the Isthmus, then took another ship to San Francisco. For the next two years young Ridgway collected in the West, returning to Washington in 1869. 632 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES In that same year, in the second paper of his budding career, Ridgway proposed that, if two populations are extremely different, even though they are connected by a chain of intermediate forms, they should be considered full species. This concept showed the effects of the morphological species definition coupled with the dawning realization that geographic variation had to be taken into consideration. One of the most able American biologists of this period was Joel Asaph Allen (b. 1838, d. 1921). In 1871 Allen demonstrated the correlation between coloration and humidity in birds, the darker populations being associated with high humidity, the lighter with aridity. He proposed that, instead of applying a name to each local population, species should be diagnosed in relation to the laws of variation. This was similar to von Gloger's (1833) suggestion. In 1877 Allen opposed the invocation of natural selection to explain the genesis of species in favor of a Lamarckian concept of the direct influence of temperature, hu- midity, food, etc. Some of these developments hardly seem like "progress" when viewed from the vantage point of 1953, but they are evidence of the problems which were under attack by the systematists of the time, whose investigations were soon to produce more durable results. In Elliot Coues (b. 1842, d. 1899) American ornithology found its genius. It was he who took the decisive step in the right direction. In 1872 in his famous Key he adopted the viewpoint that geographically complementary forms which were clearly closely related were subspecies of one species, regardless of the degree of difference between the extremes. Coues used the abbreviation "var." to indicate geographic races. The same system was employed by Baird, Brewer, and Eidgway in 1874 and remained in effect until 1881, when Ridgway took the final step to a true ternary nomenclature. There was a vast difference between the viewpoint of Schlegel and that of Coues and Ridgway. The former believed in the constancy of species and used the trinominal to designate deviations from the "type" of the species. The Americans in contrast were stanch Darwinians and for them the third name served to identify an incipient species. Baird was their mentor and he believed that if the connecting links should become extinct the previously intergrading forms would develop into distinct species. Remarkably enough there was virtual unanimity among the leaders in syste- matic ornithology in the United States regarding the concepts and usage of ternary nomenclature. Coues, the one among them with truly cosmopolitan views, decided to go to England to present the case for trinominalism and to urge the adoption of uniform rules. His optimism was not shared by Ridgway (Harris, 1928, p. 51), who felt that little would be gained. On July 1, 1884, Coues met with a group of the outstanding zoologists of England at the British Museum in South Kensington. As Ridgway had predicted, the proposals met with little enthusiasm. Only Henry Seebohm (b. 1832, d. 1895) recommended their acceptance but he was opposed by such potent adversaries as R. Bowdler Sharpe and P. L. Sclater. Coues returned home in defeat. In 1885 the Committee on Nomenclature of the American Ornithologists' Union officially accepted the concept of Coues and Ridgway. The motto of the time was "intergradation is the touchstone of trinominalism." In Europe the battle had barely begun. Seebohm (1887) had pointed out SIBLEY: ORNITHOLOGY 633 with keen insight that English ornithologists, while they accepted evolution in theory, were failing to utilize it as a working hypothesis. He urged the adoption of ternary nomenclature to differentiate nascent forms from "complete" species. Seebohm also was the first ornithologist to recognize the importance of isolation in species formation. He understood clinal variation and applied the subspecific concept solely to variation which could be defined geographically. In Germany there were some who gave trinominalism a trial but, except for Ernst Hartert (b. 1859, d. 1933), their sponsorship was carefully qualified. In 1891 Hartert went to England and in 1892 he became the director of the Tring Museum, the remarkable private museum of Walter Rothschild (b. 1868, d. 1937). After Seebohm 's death in 1895, Hartert was the only ornithologist in England consistently applying trinominals. Hartert expanded the subspecies concept to include slightly differentiated forms, even though actual intergrades were not present. Thus insular races were included in the concept and the re- quirement of intergradation was replaced with a biological interpretation of the situation in nature. For several more years the advocates of ternary nomenclature were to be looked upon as traitors but gradually they gained disciples. By 1901 Hartert was joined by several authoritative workers, the most effective being Carl E. Hellmayr (b. 1878, d. 1934). In 1903, in the introduction to his mighty work on palearctic birds, Hartert defined subspecies as geographically separated forms of the same species which are characterized not by the minor degree of the dif- ferences between them, but by differences which are related to geographical separations. The publication of Hartert's book marked the turning point, although both Sclater and Sharpe still held out. In 1909 Sharpe called the ternary system ''destructive" and gloomily predicted that all zoologists who employed it would find themselves overburdened with names. He agreed that subspecies did occur in nature but held that the binary system was sufficient for all requirements. Putting his belief into practice Sharpe raised to full species all those forms described as subspecies and thereby attained the high number of 18,939 species in his Haiid-list (5:xii, 1909). The discrepancy between that number and the recent count by Mayr and Amadon (1951) of 8,590 species is mostly due to the difference in application of the species concept. By 1912 the battle was won. The Hand-list of British Birds (Hartert, et al.) used trinominals and caused little complaint. There followed a period of re- evaluation during which many species were suppressed to the rank of subspecies and descriptions of new subspecies appeared in increasing numbers. Now that the nomenclatural practice was established it was inevitable that improvements in the causal interpretation of geographic variation and its relationship to spe- ciation would follow. As early as 1900 Otto Kleinschmidt (b. 1870) had recognized that the new concept implied that each species was composed of geographically complemen- tary forms. Kleinschmidt opposed the view that subspecies were incipient spe- cies and sought to bridge the gap between the adherents of the Linnaean species and those who believed in the nascent character of subspecies. His proposal of the term "Formenkreis," to designate a geographically complementary series of related forms was an attempt to emphasize the distinction between the new con- 634 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES cept and the Linnaean species and to overemiie tlic ohjectioiis of thoso who re- fused to give up the binary system. Kleinschmidt admitted the existence of organic evolution but believed that evolution had taken place within each Fonnenkreis following its special creation. In 1926 he elaborated upon the Formenkreis concept but the weaknesses imposed by his insistence on considering each "seed species" the result of special creation were sharply criticized. Bernhard Eensch (b. 1900) was the chief critic of Klein- schmidt's concept. Rensch (1929) clarified the concept by proposing that Har- tert's geographically varying species should be called "Rassenkreise" and two or more closely related but monotypic species which are geographically comple- mentary should be called "Artenkreise." These terms have not come into general usage but they served to call further attention to the characteristics of geo- graphically variable groups. The publication of Genetics and the Origin of Species (1937) by Theodosius Dobzhansky marked the beginning of a new phase in avian systematics. This book made a deep impression on naturalists by relating systematics to genetics. Dobzhansky was largely responsible for bringing to the attention of taxonomists the important developments in population genetics made by Sewall Wright in the United States and R. A. Fisher in England. With the realization that studies of variation were capable of producing important evidence of evolutionary processes a new method of investigation developed. In 1941 Alden H. Miller (b. 1906) forcibly demonstrated the value of examining large series of specimens in the study of variation. In his study of the avian genus Junco Miller assembled 11,776 study skins. Special trips were made to critical areas to collect adequate numbers of birds and the analysis of variation utilized statistical techniques to indicate probable as well as observable ranges of variation. Miller's Junco paper has served as the inspiration and the pattern for a number of subsequent studies of speciation undertaken by his students. Because of the relatively advanced state of avian systematics it was almost inevitable that an ornithologist would produce the first synthesized treatment of taxonomic practice and evolutionary theory. The synthesis was admirably pro- vided in 1942 by Ernst Mayr (b. 1904). In his Systematics and the Origin of Species Mayr gave systematics the first adequate integration of taxonomy, genetics, and natural history. Mayr has continued to lead in the field of evolu- tionary systematics. He was the prime mover in the founding of the Society for the Study of Evolution in 1946, and the first editor (1947-1949) of the journal Evolution. The contributors to this journal have incliided botanists, geneticists, paleontologists, and zoologists, with a wide diversity of special in- terests. The existence of the Society and the journal epitomizes the modern synthesis of fields of thought which a few years ago were regarded as diverse disciplines. Further evidence of this synthesis is provided by the recent (1953) volume on methods and principles of systematic zoology coauthored by Mayr and the entomologists, Linsley and Usinger. For a review of speciation in birds and a bibliography of recent publications see Mayr, 1950. The prediction by Sharpe in 1909 that ternary nomenclature would eventually overburden its users with names is currently finding new protagonists. The de- scription of clinal variation is difficult to accomplish with names and excessive SIBLEY: ORNITHOLOGY 635 subspecific ''splitting" has caused some systematists to propose that trinomials should be discarded or at least withdrawn from the protection of the Interna- tional Eules of Zoological Nomenclature. Thus, in 1953, w^e see the beginning of a third phase in the description of geographic variation. Just as the recogni- tion of geographic subspecies originally resulted from the combination of the development of evolutionary theory and the growth of collections so the present dissatisfaction results from the same factors. The theoretical basis has now, for continental situations at least, outstripped the descriptive method and collec- tions are now large enough frequently to reveal the true details of clinal varia- tion. It is yet too early to discern the outcome but the most hopeful approach will probably be found in a numerical evaluation of clines for only in numbers do we have a means for expressing or describing continuous variation. Although Sharpe's prediction thus proves to have contained a measure of truth, his advo- cacy of adherence to strict binomialism is certainly not the answer to the problem. The Anatomy and Classification of Birds The earliest systems of classification were based either upon external charac- ters such as the bill and foot structure or upon characteristics of habit (swim- ming, running, etc.). The famous English anatomist, Richard Owen (b. 1804, d. 1892), devoted a number of papers to avian anatomy, and the second volume of his three-volume work on vertebrate anatomy (1866-1868) was concerned to a large extent with birds. Johannes Miiller (b. 1801, d. 1858) proposed (1847) a division of the passerines upon the basis of the structure of the syrinx, a method still followed. In 1867 Thomas Henry Huxley (b. 1825, d. 1895) developed a classification of birds upon the structure and relative positions of the palatal bones. The fal- lacy of attempting to base broad conclusions upon such a narrow basis was not immediately apparent and the palatal structure has been used by many subse- quent workers as a basis for ordinal groupings. Recently (e.g., McDowell, 1948, and Hofer, 1949) there have been strong doubts cast upon the validity of Hux- ley's palatal types. Alfred Henry Garrod (b. 1846, d. 1879) fell into similar difficulties when he based his classification primarily upon the arrangement of the carotid arteries (1873a) and certain pelvic muscles (1873b, 1874). His "pelvic muscle formula" has been used extensively and George E. Hudson has recently (1937) re-evalu- ated and extended Garrod 's formula. William A. Forbes (b. 1855, d. 1883) and Hans F. Gadow (b. 1855, d. 1927) produced a long series of reports on bird anatomy. Gadow (1891) wrote the section on avian anatomy for Bronn's Klassen unci Ordnungen des Thier-Reichs. In this monumental work Gadow brought previous studies up to date and at- tempted to describe the complete morphology of the bird, including function and homologies. At about the same time (1888) there appeared the great two- volume work of Max Fiirbringer (b. 1846, d. 1920), in which he assembled an enormous amount of anatomical information and carefully weighed the charac- ters of value in classification. He recognized that the flightless groups were not necessarily monophyletic. Flirbringer's work is still the classic of bird anatomy. To some opponents of Darwinism analogy and homology were of equal taxo- 636 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES nomic rank. In 1885 E. F. von Homeyer (b. 1809, d. 1889) included the wood- peckers, nuthatches, creepers, and hoopoes in the same order, an arrangement similar to that used by Willughby more than two hundred years before! As late as 1893, Anton Reichenow (b. 1847, d. 1915) indicated his belief that a sys- tem of classification should be a means of identification and no more. The careful work of Fiirbringer and Gadow did much to overcome these viewpoints. In 1898 Frank E. Beddard (b. 1858, d. 1925), who had followed in the footsteps of Garrod and Forbes, published his volume on the structure and classification of birds, which brought together a great amount of the ana- tomical evidence for the arrangement of orders and families. Descriptive anatomy languished somewhat after the turn of the century. In the United States Robert W. Shufeldt (b. 1850, d. 1934) continued to describe the osteology of birds and in England William P. Pycraft (b. 1868, d. 1942) produced an impressive series of anatomical papers. In recent years the studies by George E. Hudson, Fred H. Glenny, and William J. Beecher have been directed toward the clarification of classification through anatomical research. Hudson has published (1937, 1948) studies on the muscles of the pelvic appendage; Glenny, beginning in 1940, has produced a series of papers on the main arteries in the region of the heart; and Beecher (1950) used the bill and jaw musculature to furnish evidence of convergent evolution in the American orioles. Other recent research has been concerned with the interpretation of functional and adaptational anatomy rather than its utilization in classification. The work of W. H. Burt (1930) on woodpecker adaptations, M. Stolpe (1932) on the hind limb, A. H. Miller (1937) on the Hawaiian goose, W. L. Engels (1940) on adaptations in the thrashers, F. Rich- ardson (1942) on tree-trunk foraging birds, H. I. Fisher (1946) on the New World vultures and William J. Beecher (1951) on the American blackbirds, are examples of this trend. Bird Migration It was within the last half of the eighteenth century that the belief that some birds hibernated in the muddy bottoms of ponds and lakes was finally discredited. With the general acceptance of the fact that birds did actually migrate there came a wave of speculation as to the methods, routes, and signifi- cance of the migratory movements. Precise data based upon observations were few at first but gradually a body of reliable information was accumulated. Among the first reliable data were records of the arrival and departure of mi- gratory species at a particular location. In 1828 Hermann Schlegel had specu- lated upon the routes and places of winter residence of European birds and the Swedish ornithologist Ekstrom had published the first arrival and departure dates of migratory species. In 1853, Karl E. Kessler, a professor at the Univer- sity of Kiev, published the arrival and departure dates for a number of species at various localities in western Russia and compared the dates with temperature. In spite of these records of actual field observation, most of the investigations into migration were conducted from a desk. J. A. Palmen (b. 1845, d. 1919), a Finnish scholar, proposed a theory of ''fly^^^ays" in 1876. He believed that there were nine narrow migratory lines which were followed by European and Asiatic SIBLEY: ORNITHOLOGY 537 birds. This concept was eventually disputed by E. F. von Homeyer (1881) who concluded that migratory birds merely followed a definite direction and that the members of a given species pass through Europe on a broad front, the width of which is equal to the width of the breeding territory. lie also stressed that the migratory direction is northeast-southwest and that the "flyways" of Palmen were the result of birds being forced together into narrow flight lines in moun- tain passes or other topographic features. Although some of his conclusions have been found in error, it was Heinrich Gatke (b. 1814, d. 1897) who gave the study of migration its greatest stimulus during the latter years of the nineteenth century. For fifty years he resided on the island of Helgoland and made observations on the hordes of migrants which paused there during the spring and fall flights. In 1891 he summarized the ideas gained from his half century of observation. Gatke agreed with von Homeyer that migration was on a broad front. He also developed the curious idea that some species which nested in Siberia reached their African wintering area by flying first west to England and then south to Africa. In the spring Gatke believed that they followed the hypotenuse of the triangle, northeast from Africa to Siberia. Recognizing the need for cooperation, Anton Reichenow and a group of col- leagues had (1875) called for help from all German ornithologists to fill gaps in the knowledge of German birds. Migratory routes were of special interest. Be- ginning in 1877 the results were published in the Journal filr Ornithologie. The practice was soon copied in England, where a committee for the study of bird migration was formed. Its first report appeared in 1879. The Ornithological Society of Vienna founded a committee on ornithological observation in 1882 and in 1883 the American Ornithologists' Union appointed the Committee on Bird Migration at the first annual meeting of the Union. C. Hart Merriam was the first chairman of the committee. These developments caused Rudolph Blasius and Gustav von Hayek of Austria to develop a plan for a world-wide network of ornithological observers. Crown Prince Rudolph of Austria commissioned them to organize the First In- ternational Ornithological Congress, which met in Vienna in 1884. Blasius be- came the chairman of a committee to organize the observers of the world and the publication Ornis was founded and first published in 1885. The undertaking did not succeed long. The mass of uncritically accepted data was of greatly variable value and no one was willing to undertake its analysis. By 1890 the various branches had again become autonomous. In America the Committee on Bird Migration had enthusiastically set to work. Merriam's energy and knowledge had combined to push the project along. By 1885 the job had grown too large for the American Ornithologists' Union and in 1886, when Merriam became head of the Division of Economic Orni- thology and Mammalogy, the migration studies were continued under govern- mental auspices. In 1888 Wells W. Cooke (b. 1858, d. 1916), who eventually became the "bird migration expert" of the Bureau of Biological Survey, pub- lished his classic report on migration in the Mississippi Valley. This paper at- tempted to correlate weather data with observations on migration and marked the beginning of such investigations in North America. As valuable and important as these studies were, they were limited by the 638 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES available data. Phenology did not provide information concerning the speed and direction of individual migrants; a new technique was needed. Individual birds had been marked on many previous occasions but the at- tempts were sporadic and of short duration. It was a Danish schoolmaster, Hans Christian Cornelius Mortensen (b. 1856, d. 1921) v/ho first attempted to band birds in a systematic fashion. His first trial in 1890 was with zinc bands in- scribed "Viborg 1890." Two such bands were placed upon starlings. Mortensen noted that the bands seemed to be unpleasant to the birds and gave up the project. A few years later aluminum came into use for poultry bands. A mer- ganser which Mortensen banded with one of these was shot soon after and the band was returned to him. In 1899 he captured and banded 162 adult starlings but no returns were received. The experiment was repeated in 1900 with bands stamped "M. Danmark" and this time his banded birds were shot in Holland and Norway. The technique had proved successful. Banding developed rapidly. In 1900 the German Ornithological Society (Deutschen Ornithologen-Gesellschaft) subsidized and founded the now famous "Vogelwarte Rossitten." This bird observation station located at the town of Rossitten on the narrow coastal spit of the Kurische Nehrung was placed under the direction of Johannes Thienemann (b. 1863, d. 1938). The principal objec- tive was the study of migration. Banding was begun in 1903, using aluminum rings carrying a number and the year. By 1937 over 763,000 birds had been banded at Rossitten and returns totaled more than 10,000. The Rossitten sta- tion remained active until World War II and produced a large number of signifi- cant papers. The idea of banding spread rapidly and Avas adopted by other organizations and individuals. Paul Bartsch (b. 1871) of the United States Na- tional Museum banded 101 fledgling black-crowned night herons near Washing- ton, D. C, in 1902 and 1903, and in 1902 Leon J. Cole (b. 1877, d. 1948) pro- posed the systematic use of banding as a means of studying migration. Other investigators soon followed these pioneers and by 1909 banding had be- come important enough to siiggest the need for an organized effort. The Ameri- can Bird Banding Association was formed in New York on November 8, 1909. For the next decade the work was sponsored by various organizations, including the Linnaean Society of New York and the New Haven Bird Club, in addition to the American Bird Banding Association. In 1920 the Bureau of Biological Survey took over the responsibility of furnishing bands and maintaining the records and Frederick C. Lincoln was placed in charge of the project. Over 1,000,000 birds had been banded in the United States and Canada by 1933 and nearly 6,000,000 by 1949. The year 1909 also saw the formation of two banding organizations in Great Britain. A. Landsborough Thomson founded the Aberdeen University Bird- Migration Inquiry and Mr. H. F. Witherby launched a banding program in con- nection with the magazine British Birds. In 1937 the latter program was trans- ferred to the control of the British Trust for Ornithology, with headquarters in the British Museum (Natural History). By 1927 there were seven European countries operating banding stations. In 1950 Rydzewski listed banding stations in eighteen European countries, Egypt, South Africa, India, Japan, Australia, New Zealand, Canada, and the United States. The German bird observation stations of Rossitten and Helgoland have moved SIBLEY: ORNITHOLOGY 639 to new locations since AVorld War II. The Rossitten group, under Ernst Schiiz, is now at Radolfzell on Lake Constance, while the Helgoland station has moved to Wilhelmshavn and is directed by Rudolf Drost. The first bird observation sta- tion in Sweden was founded at Ottenby in 1945 and others (e.g., Fair Isle, Isle of May, Skokholm) have been intermittently active in the British Isles, The information derived from banding includes much in addition to data concerning migratory routes. Knowledge of the dispersal of juveniles, sex ratios, speed of flight during migration, longevit}^, plumage change in relation to age, and diseases and parasites are among the items to which banding has made a direct or indirect contribution. Thanks to the banding technique the mysteries of bird migration were fewer in 1920 than they had been in 1900, but at least two major problems remained unsolved. AVhat was the stimulus which started a bird off on its migratory flight with such remarkable precision, and how did the migrating bird find its way? These questions demanded experimental investigation. The precision with which migratory birds arrived at a given point year after year was proof that the timing device which provided the stimulus was equally precise. The annual cycle of weather, seasonal variation in food supply, and other phenomena had been suggested as the source of the stimulus. These w^ere too variable to account for the regularity of migration. When Professor AVilliam Rowan of the Univer- sity of Alberta began his investigation into the problem, he had logically settled upon the annual cycle of changing day-length as the only apparent cyclic phe- nomenon with the necessary degree of precision. This hypothesis he set out to test. In the fall of 1924 Rowan trapped southbound slate-colored juncos pass- ing through Edmonton. The birds were caged in outdoor aviaries, one of which contained an electric light. The experimental procedure was classically simple. Beginning on November 1 the light in the experimental cage was left burning for 71/2 minutes after dark. A daily increment of 71/2 minutes was added until December 3, when the increment was reduced to 5 minutes. By December 15 this procedure resulted in the light remaining on until 11:00 p. m. The increases were then discontinued, the light going out at 11:00 p. m. until January 9, when the experiment was terminated. Although their environment was that of a Canadian winter the gonads of the experimental birds had attained the maxi- mum breeding size, and the males were in full song. The gonads of the control birds in the unlighted cage were at the winter minimum size. Here indeed was proof of the effect of photoperiodism on the sexual cycle. Rowan's little book The Riddle of Migration (1931) summarizes his experiments. Although some of Rowan's conclusions concerning the relationship between the gonad cycle and the migratory impulse have been modified, his experiment started the inten- sive investigations, such as those of Wolf son (1945), which have led to an under- standing of the annual stimulus for migration. The present state of knowledge has been summarized by Earner (1950), who proposes a working hypothesis which attempts to reconcile many seemingly divergent facts and suppositions. This hypothesis, somewhat simplified, states that twice each year migratory spe- cies of birds come into a distinct physiological condition which places the bird in a "disposition to migrate." This is indicated by the deposition of fat in many species. The gonads begin to increase in size and a condition of "restlessness," which is especially noticeable in caged birds, is evident. 640 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES The fundamental cycle which periodically places the bird in the "disposition to migrate" is probably the result of the cyclic function of the anterior lobe of the pituitary gland. This cycle could be, and probably is, the result of periodic change in stimulation by periodic external environmental factors such as day- length. When a certain threshold of stimulation is reached, the bird is stimu- lated to migrate. The act of migration, that is, the actual movement through space on a remarkably exact time schedule, is the result of a complex inherited behavior pattern stereotyped in the nervous and endocrine systems. The problem of orientation and navigation during the migratory flight posed a still more difficult problem. The classical experiments of J. B. Watson and K. S. Lashley (1915) had proved that nesting birds could find their way "home" over long, unfamiliar routes. Further experiments on homing have been car- ried out by Riippel in Germany, Lockley in England, and Griffin in the United States. All tended to confirm the fact of homing ability in birds but failed to yield unquestionable proof of the method of orientation. The importance of landmarks in the homing of carrier pigeons was established by several workers, including the Heinroths (1941). The hypotheses presented by Ising (1945) and Yeagley (1947), postulated that orientation could be achieved by detection of variations in the fields of force resulting from the earth's rotation (Coriolis force) were vigorously attacked by both physicists and biologists (see Odum, 1948). The most promising development in the field of orientation research is the work of Gustav Kramer (1949, 1950), who has successfully demonstrated that the sun is utilized in orientation at least by certain diurnal migrants. Kramer constructed a round cage having six equally spaced windows. Each window was equipped with a hinged shutter upon which a mirror was mounted. By manipu- lation of the shutters the angle of the sun's rays entering the cage could be modi- fied. With the shutters wide open a spring migrant European starling {Sturnus vulgaris) made repeated attempts to fly toward the northwest, the normal di- rection for the spring migration. AVhen the mirrors were placed so as to deflect the direction of the incidental light by 90° the captive bird changed the direc- tion of its flight in accordance with the direction of the light. There still remains the problem of orientation by nocturnal migrants but Kramer's experiment will certainly direct further research along profitable pathways. The paper by Drost (1950) provides a review of much of the recent work on bird migration. Bird Behavior The necessity of objectivity as a component of the scientific method is unde- niable. It is equally certain that no field of endeavor has had a more difficult time incorporating the objective viewpoint into its investigations than that of animal behavior. Not until it emancipated itself from the burden of anthropo- morphism was it able to attack its problems with any measure of success. The viewpoints of Christian Ludwig Brehm (b. 1787, d. 1864) and his son Alfred Edmund Brehm (b. 1829, d. 1884) dominated the thinking on bird be- havior during the mid-nineteenth century. The younger Brehm's two great works, Das Lehen der Vogel (1861) and IHustrirtes Thierlehen (1864-1869), SIBLEY: ORNITHOLOGY 641 went through several editions and were translated in various degrees into other languages. Brehm's viewpoint was strongly anthropomorphic and sentimental and, since his influence was great, he was accepted as authoritative. The first serious challenge to Brehm's views was presented by Bernard Altum (b. 1824, d. 1900) in his now famous classic, Der Vogel und Sein Lehen (1868). Altum 's viewpoint was anti-Darwinian but also anti-anthropomorphic. He proposed a strongly instinctive mode of behavior for birds and believed that their activities were the result of unthinking reactions to external stimuli. Altum 's fame is secure as the first to expound the concept of territory in birds, if for no other reason. His discussion of territorial behavior includes an analysis of the function of song as a threat to other males and an invitation to females and the importance of territory in reducing competition for food between mem- bers of a species. (For a review of Altum's territorial concept see Mayr, 1935.) The reaction to Altum's ideas was immediate and mostly hostile. Brehm's influence was so great that it was twenty-five years before Altum's views were generally accepted in Germany. Despite the seemingly revolutionary and advanced concepts expressed by Altum he did not attract much attention outside of Germany and even there the importance of his ideas was not fully realized. This situation was probably due to the general lack of interest in psychological problems among ornitholo- gists during the latter part of the nineteenth century. Progress came slowly, and for some time it was not due to the work of orni- thologists but to the investigations of psychologists and general biologists. The work of C. Lloyd Morgan (b. 1852, d. 1936) was unnoticed by most ornitholo- gists but gradually his ideas concerning instinctive behavior became known to a few. Morgan's theory of instinctive behavior (1896) was basically mechanistic and was founded in Darwinism. He believed that instincts are innate and that they become fixed by selection. He also found reason to believe that an instinc- tive chain of acts could be modified through the conscious activity of the animal and he called this type of modification an "acquired instinct." According to Morgan, an animal inherited a basic set of instincts but was able to learn by experience and add to its innate instinctive behavior. As the concepts of psychology developed, a number of ornithologists began to apply them to the study of living birds. In the United States Francis H. Her- rick (1901) was among the first to utilize photography and careful observation of the living bird in studying behavior. He emphasized that nest-building and other avian activities are purely instinctive acts in which the bird exhibits no power of choice. Arthur A. Allen's study (1914) of the red-winged blackbird (Agelaius plioeniceus) was an important milestone in avian ecological and life history research which influenced and set the pattern for numerous studies of single species by his students and other workers. In England the study of the living bird found an able and articulate protagonist in Edmund Selous (b. 1858, d. 1934). His books, Bird Watching (1901), Bird Life Glimpses (1905), and The Bird Watcher in the Shetlands (1905b), promoted the value of the note- book and binocular as tools of ornithology. His philosophy was amusingly pre- sented in verse when he wrote : Some men have strange ambitions. I have one: To make a naturalist without a gun. 642 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Selous correctly pointed out that there was little information available on birds except about their plumages, nests, eggs, and distribution. He stressed the need for studies of behavior and the need for interpretation of habits, in addition to mere factual recitation. His viewpoint was Darwinian and his interpretations of behavior were in terms of selection and survival. Two years later, H. Eliot Howard (b. 1873, d. 1940) began his studies of the British warblers (1907-1915). Howard was concerned primarily with the investigation of nesting and brood care. His study of the several species of Brit- ish warblers permitted comparison of the breeding biology of a group of closely related species. In 1910 Howard called attention to the fact that the males in many species took up a territory and defended it against intruders. He de- veloped the territorial concept, apparently unaware of the work of Altum (1868), and in 1920 devoted an entire volume. Territory in Bird Life, to the subject. Howard's carefully documented theory had an immense effect on the ornitholo- gists of the entire world. Others had recognized the general facts and had even stated the elements of the theory of territory but Howard's emphatic presenta- tion became the starting point of a new era in the study of avian behavior. Among the many important studies which have utilized and expanded the ter- ritorial concept since 1920 are two which rank as classics. As a result of several years of intensive work on the song sparrow {Melospiza melodia) near Colum- bus, Ohio, Margaret Morse Nice published two volumes (1937, 1943) dealing with its life history and behavior. Mrs. Nice's familiarity with the literature of avian behavior permitted a truly comparative presentation and her methods have served as the model for numerous subsequent investigations. In 1941 Mrs. Nice prepared a valuable review of the territorial concept which includes a com- prehensive bibliography of the subject. The well-founded tradition of field natural history characteristic of present- day England, begun by Gilbert White (1789) and nurtured by Edmund Selous, is today led by David Lack. Lack's fine study (1939) of the life history of the English robin {Erithacus rubecula) ntilized the techniques of observation of marked individuals (color-banded) and the experimental use of stuffed speci- mens. His work served to focus attention on the value of the intensive study of single species. The first to bridge the gap between behavior and systematics was Oscar Hein- rotli (b. 1871, d. 1945), who presented the idea (1910) that voice and behavior were clues to relationship. Heinroth's interest was in the living bird and many of his behavior studies were on captive birds in the Berlin Zoo. Heinroth laid the foundation for further research in comparative behavior and crowned his life's work with the remarkably detailed three-volume work, Die Vogel 3Iittel- europas (1924-1928), with his wife as coauthor. This ambitious project was in preparation for twenty years and included nearly three thousand photographs and descriptions of the details of behavior, development, and other phases of life history. The study of instinctive behavior received a new impetus with the work of Konrad Lorenz in the 1930's. At his home in Altenberg, Austria, Lorenz studied free-living, semitame birds of several species. His work on the behavior of the jackdaw {Corvus monedula) started in 1925 with a single bird. A flock was gradually built up which provided research material for a number of ethologi- SIBLEY: ORNITHOLOGY 643 cal studies (e.g., 1931). In 1935 Lorenz proposed the "releaser" concept to ex- plain the initiation of instinctive behavior patterns. In an English version of the 1935 paper Lorenz (1937, p. 249) defined a "releaser" as follows. The means evolved for the sending out of key-stimuli may lie in a bodily character, as a special color design or structure, or in an instinctive action, such as posturing, "dance" movements and the like. In most cases they are to be found in both, that is, in some instinctive acts which display color schemes or structures that were evolved exclu- sively for this end. All such devices for the issuing of releasing stimuli, I have termed releasers (Ausloser) , regardless of whether the releasing factor be optical or acoustical, whether an act, a structure or a color. The releaser concept found general acceptance among students of behavior and was quickly applied to other studies. It was the unifying principle which had been lacking and which greatly simplified much of the complicated termi- nology that had enmeshed the study of animal behavior. Owing largely to Lo- renz the problem of innate behavior has received a great deal of attention in the past fifteen years. There have been many ethological studies utilizing the "releaser" concept. The principal contributor has been Nikolas Tinbergen, formerly of the Uni- versity of Leiden, now Lecturer in Animal Behavior at Oxford. Tinbergen has successfully developed the objectivistic approach to the analysis of animal be- havior. His work has included study of the orientation mechanism of the digger wasp (Philanthus) , territory and breeding behavior of the three-spined stickle- back {Gasterosteus aculeatus) , and numerous investigations of avian behavior. Among the latter his study (1939a) of the spring behavior of the snow bunting {Plectrophenax nivalis), and the analysis of the releaser for the begging re- sponse in herring gull (Larus argentatiis) chicks (with H. C. Perdeck, 1950) are examples. Tinbergen's ability to synthesize has been of great value to other ornithologists. His extensive knowledge of this complex field has made possible several valuable "review" papers (1936, 1939b, 1942, 1948) and recently (1951) has resulted in a book which summarizes the present state of knowledge of in- stinctive behavior. Fossil Birds The history of paleornithology is nearly coterminal with the span of our Century of Progress. Few discoveries of importance were made before 1861, when the remains of Archeopteryx were found in the lithographic limestone quarry at Solenhofen, Bavaria. The skeleton of this Upper Jurassic link be- tween reptiles and modern birds was described by Owen in 1863. In 1877 a second Jurassic bird was found near Eichstatt, Bavaria. It was described by Dames in 1884 as Archeopteryx siemensi. In 1921 Petronievics made this second fossil the type of the genus Archeornis. Both specimens combine numerous rep- tilian characters with the presence of feathers. There is general agreement that these Jurassic fossils represent the first birds although Lowe (1944) believes that they should be considered flying reptiles. In the preparation of his four volumes on the fossil birds of France (1867- 1871) Alphonse Milne-Edwards (b. 1835, d. 1900) visited all the large geological collections in Europe. He assembled more than 4,000 fossil bones and the skele- 644 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES tons of nearly 800 species of living birds for comparison. The illustrations are accurate enough to serve as the basis of comparison by critical modern workers. The first important contribution to the study of fossil birds in America was made in 1870 by 0. C. Marsh (b. 1831, d. 1899). In 1872 Marsh announced the discovery of the Cretaceous toothed bird, Hesperornis regalis. Marsh continued to describe avian fossils and in 1880 published a monograph on the Cretaceous toothed birds of North America. Marsh described a total of 40 species of fossil birds during his lifetime. Edward Drinker Cope (b. 1840, d. 1897), Marsh's famous rival, described his first avian fossils in 1871 and the giant Eocene Diatryma from New Mexico in 1876. One of the most prolific writers on osteology and paleornithology was Robert W. Shufeldt (b. 1850, d. 1934). In 1891 he began his descriptions of fossil birds, which were to number 43 species, more than the total of any other North Ameri- can worker to date. In South America, Florentino Ameghino (b. 1854, d. 1911) described (1891) the gigantic flightless ]\Iiocene bird Phororhacos, which stood at least seven feet tall and had an enormous raptorial beak. In 1895 Ameghino's book on the fossil birds of Patagonia appeared. From the lower Eocene beds near Croyden, England, E. T. Newton described (1886) a huge flightless bird, Gastornis, larger than an ostrich, which may be related to the ducks and geese (Swinton, 1934). Most of the avian fossils discovered before 1909 were those of large, flightless species. This is not surprising, for flying birds are less likely to become ensnared in natural traps and the bones of small birds are so fragile as to reduce the chances of intact preservation in ordinary sediments. In 1909 Loye Holmes Miller began the study of the abundant Pleistocene material preserved in the asphalt traps of Rancho La Brea, McKittrick, and Carpenteria in southern California. Among the numerous bones of large raptors and scavengers were thousands of skeletal elements belonging to small passerines. As a result of the studies by Loye Miller, and later by Hildegarde Howard and Alden Miller, the Pleistocene avifauna of California is the most completely known fossil avifauna in the world. From Rancho La Brea alone 105 species have been identified. Since 1920 the most active paleornithologist in North America has been Alex- ander Wetmore. In 1921 he described an owl from the Eocene of Wyoming and has since described a number of Tertiary birds, primarily from the Miocene and Pliocene. His check-list (1940) of the fossil birds of North America includes 165 forms which are still living and 184 extinct species. This list has increased but slightly since 1940. Two valuable references to avian fossils have appeared in recent years. In 1926 Gerhard Heilmann's The Origin of Birds presented the results of his studies on the relationships between reptiles and birds. Heilmann amassed anatomical and embryological evidence to support the idea of the reptilian origin of birds. His book contains valuable and detailed studies on Archeopteryx and xircheornis. The Handbuch der Palaeornitkologie (1933) by Kalman Lambrecht provides a review of the world-wide knowledge of fossil birds. One consequence of the development of knowledge of fossil birds has been speculation as to the origin of flight. Marsh (1880) suggested a tree-dwelling SIBLEY: ORNITHOLOGY 645 ancestor while Nopcsa (1907) derived flying birds from rapid-running ground- dwelling forms. Beebe (1915) proposed the "tetrapteryx" stage as an ancestral intermediate form. This hypothetical progenitor had a ''pelvic wing" which Beebe believed to be indicated by the femoral tract of modern birds. Steiner (1917) proposed a proavian which is both tree-dwelling and running, with long hind limbs and forelimbs equipped with functional claws and an expanded air-foil of feathers. Two recent papers by Hildegarde Howard (1947, 1950) present evidence of avian evolutionary history based on the fossil record while AVetmore (1950) has reviewed the addition to the knowledge of fossil birds since the publication of Lambrecht's book in 1933. Ornithological Periodicals It is doubtful if any other class of animals has been the inspiration for the founding of as many serial publications as birds. Most of these journals have enjoyed but a brief life and few have become scientifically important. A small number have been privately printed; most have been or are the organs of so- cieties or institutions. Within the pages of the Journal filr Ornithologie, the Ihis, and the Auk have appeared more than half the basically important ornithological papers of the past century. These three have enjoyed the benefits of an active membership in the supporting societies and that all-important necessity, good editorship over long periods of time. It was partly as a protest against the provincialism of other ornithological periodicals that Gustav Hartlaub (b. 1814, d. 1900) and Jean Cabanis (b. 1816, d. 1906) founded the Journal filr OrnitJioJogie in 1852. With Cabanis as editor and leading German ornithologists as contributors the "J. f. 0." soon became the principal German ornithological periodical. The present editor is Erwin Strese- mann. In 1858 the Ihis was founded in England as the organ of the British Orni- thologists' Union. It too enjoyed a series of competent editors and quickly be- came the premier ornithological periodical in English. Among its editors have been Alfred Newton, Osbert Salvin, Philip Lutley Sclater, and his son, William Lutley Sclater. The present editor is R. E. Moreau. The Nuttall Ornithological Club was organized in Cambridge, Massachusetts, in 1876 and began publication of The Bulletin of the Nuttall Ornithological Club in the same year. Seven years later, when the American Ornithologists' Union was organized in New York (September 26, 1883), the Nuttall Club of- fered its Bulletin and its editor as the foundation for the journal of the union. The Auk was chosen as the name of the new journal and J. A. Allen continued as editor until 1912, when he was succeeded by AVitmer Stone. Glover Morrill Allen followed Stone in 1937. Following in succession as editor were John T. Zimmer, Harvey I. Fisher and Robert AV. Storer (incumbent). The official date of the founding of the Wilson Ornithological Club is Decem- ber 3, 1888, although its roots go back to 1858 under various names. In 1889 a journal was started, the Ornithologists and Oologists' Semi-Annual. Within the next nine years the name was changed no less than six times, the seventh (1898), 646 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES being The Wilso7i Bulletin, which survives today. At first devoted primarily to the field ornithology of the jMiddle West it has more recently included papers of wide scope and high quality. Among its editors have been Lynds Jones, Thomas C. Stephens, and Josselyn Van Tyne. The Cooper Ornithological Club was organized on June 22, 1893. The Bulle- tin of the Cooper Ornithological Cluh began publication in 1899; in the follow- ing year the name of the journal was changed to The Condor. As editor from 1906 to 1939, Joseph Grinnell was largely responsible for its continuing success. His high standards have been continued by Alden H. Miller. In 1952 the name of the organization was officially changed to Cooper Ornithological Society. The following list includes a world-wide representation of the periodicals de- voted entirely to ornithology. Aquila, founded 1894 in Hungary. Printed in both Hungarian and German. The Emu, founded in 1900 as the official organ of the Australasian Ornithologists Union. (Australia.) Britisli Birds, founded in 1907. Devoted primarily to the occurrence and behavior of the birds of Great Britain. Tori, founded in 1915 as the bulletin of the Ornithological Society of Japan. In Japanese. El Hornero. founded in 1917 by the Ornithological Society of La Plata (Argentina). The principal ornithological journal of South America. In Spanish. UOiseau, founded in 1920, and Alauda (1929) are the principal periodicals of France. Le Gerfaut (1909-1914, 1919-), published by the Belgian Central Ornithological So ciety. In French. The Ostrich (1930), journal of the South African Ornithological Society. Austin Rob- erts was the first editor. Bird-Banding (1930), published by the Northeastern Bird-Banding Association (New England region). Includes reviews of the literature of avian biology. De7- Vogelsug (1930-1943), devoted primarily to studies of bird migration and pub- lished by the German bird observation station at Rossitten (Vogelwarte Rossitten). Publication suspended in 1943 during World War II. In 1948 the publication Die Yogel- ivarte replaced Vogelzug as the organ of the German bird observation stations. Ornis Fennica (1924), published by the Ornithological Society of Finland. Papers in Finnish, German, or Swedish. ArcZea (1912), published by the Netherlands Ornithological Society. Limosa, founded in 1928 as the Orgaan der Cluh van Nederlandsche Vogelkundigen. Became Limosa in 1937. The preceding list includes some of the more enduring and important peri- odicals which contain only ornithological papers. In addition, many papers deal- ing with birds regularly appear in such journals as the Proceedings of the Zoological Society of London, Evolution, and the journals of psychology, physi- ology, anatomy, etc., in all languages. The "occasional papers," "proceedings," "transactions," "novitates," "comp- tes rendus," "archives," etc., of museums and universities are other important sources of ornithological literature. The Zoological Record is undoubtedly the most nearly complete bibliographic reference source for zoological literature. The "Aves" section averages nearly 1,500 references per year. It is certain that well over 100,000 books and papers on birds have been published during our Century of Progress. sibley: ornithology 647 Ornithological Monographs The results of most original research today are customarily published in the periodical journals. This material is often widely scattered and frequently unavailable or unknown to many interested persons. Fortunately, it is also cus- tomary for specialists to syntliesize the numerous research papers and to pro- duce books which summarize their fields of endeavor. In ornithology there are innumerable books dealing with the distribution and occurrence of the birds of areas ranging in size from a university campus to the world itself. These vary from mere lists to extensive compendia containing enormous amounts of in- formation. There have been far fewer books devoted to such subjects as be- havior, anatomy, and other phases of avian biology. Books which fall into these categories number in the thousands. Rather than try to cite numerous examples and thereby omit reference to many equally worthy of inclusion, it seems better to single out a few major works published before 1900 and to give more space to the important volumes of the past fifty years. Volumes which have been noted elsewhere in this chapter will not usually again be cited. The general faunistic works on European birds are seemingly endless. The British Isles have been especially prolific of local faunal compilations. Follow- ing in the footsteps of William Yarrell (b. 1784, d. 1856), whose History of British Birds (1837-1843) was long the standard, was Howard Saunders (b. 1835, d. 1907), who brought Yarrell's work up to date in 1889 and further re- vised it in 1899. Today the standard work is the five-volume Handbook of H. F. Witherby and his collaborators (rev. ed., 1943). This remarkable compilation has no parallel in English but is in some ways comparable to the work of the Heinroths (1926-1928) on central European birds. Germany, too, has pro- duced a spate of faunal treatises. Niethammer's recent (1937-1942) three- volume handbook is outstanding. For Europe in general there is the magnificant nine-volume treatise by Dres- ser (1871-1890) and Hartert's (1903-1923) scholarly three volumes on pale- arctic birds. African birds have been the subject of numerous books. Hartlaub (1857), Finsch and Hartlaub (1870), Shelley (1896-1912), Bannerman (1930-1951), and Chapin (1932-1939) are among the many contributors. The English extended their interest in natural history to all parts of the British Empire. The ornithological volumes of The Fauna of British India (1889-1898) were prepared by E. W. Gates and W. T. Blanford. In 1922 E. C. Stuart Baker published the first volume of a revised edition of this work. Australian birds were first extensively described in a monograph by John Gould (b. 1804, d. 1881), whose seven volumes (1840-1848) were illustrated with 600 hand-colored plates and followed by a supplement containing 81 more (1851- 1869). In spite of his unfortunate prolixity for generic splitting the work of Gregory H. Mathews (b. 1870, d. 1949) is pre-eminent in Australian ornithology. His twelve large volumes (1910-1928) are among the last of the elaborately il- lustrated extensive faunal monographs. The birds of New Zealand were treated by Walter L. Buller (b. 1838, d. 1906) in 1872-1873 and more recently (1930) by W. R. B. Oliver. 648 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES The avifaunas of most Asiatic countries have been treated in monographs. Many of these are listed by Casey Wood in his Introduction to the Literature of Vertebrate Zoology (1931, pp. 77-78). Among the numerous faunal works on New AVorkI birds is the detailed sys- tematic treatment of the birds of North and Middle America (1901-1950) by Eobert Ridgway (b. 1850, d. 1929), which has been continued since Ridgway's death by Herbert Friedmann. In 1918, Charles B. Cory (b. 1857, d. 1921) be- gan the publication of an extensive catalogue of all of the species and subspecies of the Americas and adjacent islands. This work was continued by Charles E. Hellmayr (b. 1878, d. 1944) after Cory had completed two volumes. The last four of the fifteen volumes were finally finished by H. B. Conover from Hell- mayr's manuscript. The ''Aves" volumes of the Biologia Centrali- Americana (1879-1904) by Sal- vin and Godman described over 1,400 species of Central American birds. The authors, opponents of trinominal nomenclature, maintained a consistently bi- nominal treatment in their work. Among more recent systematic treatments of Central American birds is that of Dickey and van Rossem on El Salvador (1938) . The books by Sclater and W. H. Hudson (1888-1889) and by W. H. Hudson (1920) on Argentine birds are among the best known of many volumes on South American birds. Among recently active workers have been John T. Zimmer on Peruvian birds (1931, et seq.) and AVilliam H. Phelps and William H. Phelps, Jr., mainly on the birds of Venezuela. To date there has been only one attempt to describe all of the known species of birds in the world. It was the indefatigable Richard Bowdler Sharpe (b. 1847, d. 1909) who set this as his task shortly after he succeeded G. R. Gray as keeper of the bird collection of the British Museum (Natural History) in 1872. The first volume of the Catalogue of the Birds in the British Museum ap- peared in 1874. Of its twenty-seven volumes Sharpe himself wrote fourteen. Among others who contributed volumes to this remarkable undertaking were P. L. Sclater, G. E. Shelley, T. A. Salvadori, 0. Salvin, and E. Hartert. These volumes include plumage descriptions, synonyms, references, and distributional information. Sharpe's Hand-list (1899-1909) was also the first world-wide check-list. In 1931, James Lee Peters (b. 1889, d. 1952) published the first volume of his Check-list of Birds of the World, seven volumes of which had been completed by 1951. In terms of numbers of species this is approximately the halfway point. Elaborately illustrated monographs of genera, families, or orders were pro- duced in numbers during the latter part of the nineteenth century. John Gould (b. 1804, d. 1881) wrote and illustrated a number of famous works of this na- ture including the hummingbirds (1849-1861), which occupied five volumes and contained 360 colored plates. Otto Finsch (b. 1839, d. 1917) wrote a monograph on the parrots of the world (1867-1868), which is still the most complete ac- count of the group. In the United States, Daniel Giraud Elliot (b. 1835, d. 1915) has published monographs on the grouse (1864-1865), the pheasants (1870-1872), the hornbills (1877-1882), the North American shore-birds (1895), and several other groups. More recent monographic treatment has been accorded the pheasants by Beebe (1918-1922) and by Delacour (1951), the birds 9f prey by Sw^ann and SIBLEY: ORNITHOLOGY - 649 Wetmore (1924-1945), and the clucks by Phillips (1922-1926). Murphy's two volumes on the oceanic birds of South America (1936) include extensive mate- rial on life history and behavior in addition to taxonomic and distributional data. The thick volume by Knowlton (1909) is the only attempt to date to provide a survey of the habits, appearance, and distribution of the birds of the entire world. All families are considered with attention given to significant species in each. Newton's Dictionary of Birds (1893-1896) is an alphabetically ar- ranged compendium of various phases of ornithology. In marked contrast to the enormous number of volumes dealing with faunal or systematic groups is the paucity of works on avian biology. This situation is partly due to the fact that systematics must precede studies of the living animal and it is only recently that the classification of birds has attained the necessary degree of completeness. A second factor is the relative novelty of the basic concepts upon which interpretations of behavior, physiology, etc., are founded. Among the first books which tried to bring together the information on bird biology were those of Beebe (1906) and Pycraft (1910). In 1923, J. A. Thom- son published his volume on bird biology, which included chapters on adapta- tion, behavior, migration, and so forth. The major work on avian biology to date was written by the dean of world ornithologists, Erwin Stresemann (b. 1889) and published (1927-1934) as a volume of Kiikenthal and Krumbach's Handbuch der ZooJogie. This moniunental book contains extensive discussions of anatomy, physiology, and other phases of avian biology. Before Stresemann 's volume was completed, there appeared the first parts of Franz Groebbel's (b. 1888) detailed treatment of avian anatomy and biology (1932-1937). In 1950 a collaborative effort by a group of twelve French biologists under the direction of Pierre Grasse produced a volume which, while variable in the extent and quality of the treatment of its different sections, is the onh^ readily available up-to-date compendium on the biology of birds. It contains chapters on anatomy, physiology, genetics, behavior, embryology, ecology, etc., and a sys- tematic synopsis of the birds of the world. Of importance to students of avian biology are such volumes as Friedmann's studies on social parasitism in the cowbirds (1929) and the parasitic cuckoos of Africa (1948), the compendium by Armstrong (1947) on bird behavior and N. Tinbergen's recent (1951) book on instinct. The book on bird parasites by Miriam Rothschild and Theresa Clay (1952) brings together for the first time the large and scattered literature on this subject. Anyone familiar with the literature of ornithology will think of numerous works, as important as some herein included, which have been omitted. The attempt has been to select examples, not to survey the entire literature of the past century. It has not been possible in this brief survey of ornithology during the past century to cover all of the aspects of the subject. Omission of such important phases of research as bird flight, avian genetics, ecology, endocrinology, and other subjects is regretted. For the reader interested in further historical in- 650 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES formation there is the recent and scholarly volume by Erwin Stresemann (1951), which has provided the foundation for several sections of this chapter. The debt to Professor Stresemann is gratefully acknowledged. The little volume by Maurice Boubier (1925) was also useful, as were the chapters in Fifty Years' Progress of American Ornithology, 1883-1933 published by the American Orni- thologists' Union on its fiftieth anniversay. Casey Wood's (1931) survey of the vertebrate literature and R. M. Strong's (1939-1946) bibliography were re- peatedly consulted. To my colleague and friend, Dr. William Graf, I owe a debt of gratitude for his patient and extensive help with the translation of large portions of Dr. Stresemann 's book. LITERATURE CITED Allen, A. A. 1914. The Red-winged Blackbird: A study in the ecology of a Cat-tail marsh. Proc. Linn. Soc. New York, No. 24. Allen, J. A. 1871. On the mammals and winter birds of East Florida, with an examination of certain assumed specific characters in birds, and a sketch of the bird-fauna of eastern North America. Bull. Mus. Comp. Zool., 2:161-450. 1877. The influence of physical conditions in the genesis of species. Radical Review, 1:108-140; reprinted in Ann. Rept. Smithson. Inst., 1905:375-402. Altum, B. 1868. Der Vogel und sein Leben. 240 pp. Munster. Ameghino, F. 1891. Enumeracion de las aves fosiles de la Republica Argentina. Rev. Argentino Hist. Nat., 1:441-456. 1895. Sur les oiseaux fossiles de Patagonie. Bol. Inst. Geog. Argentino, 15:501-602. Armstrong, E. A. 1947. Bird Display and Behaviour, iii + 431 pp. London: Lindsay Drummond. Baird, S. F. 1859. Notes on a collection of birds made by Mr. John Xantus, at Cape San Lucas, Lower California, and now in the museum of the Smithsonian Institution. Proc. Acad. Nat. Sci. Phila., 11:299-306. Baird, S. F., T. M. Brewer, and R. Ridgway 1874. A History of North American Birds. 3 vols. Boston: L. H. Brown and Co. Baird, S. F., J. Cassin, and G. N. Lawrence 1858. Pacific Railroad Reports. Volume IX, Birds. Ivi + 1005 pp. Washington, D. C. Baker, E. C. S. 1922-1930. The Fauna of British India, including Ceylon and Burma. (Birds.) 8 vols. London: Taylor and Francis. Bannebman, D. a. 1930-1951. The Birds of Tropical West Africa. 8 vols. London: Crown Agents for Colonies. Bartsch, p. 1903. Notes on the herons of the District of Columbia. Smithson. Misc. Coll., 45: 104-111. SIBLEY: ORNITHOLOGY 651 Beddard, F. E. 1898. The Structure and Classification of Birds, xx + 548 pp. London and New Yorlt: Longmans, Green and Co. Beebe, C. W. 1906. The Bird, its Form and Function, xii + 496 pp. New Yorli. 1915. A Tetrapteryx stage in the ancestry of birds. Zoologica, 2:39-52. 1918-1922. A Monograph of the Pheasants. 4 vols. London: New York Zoological Society. Beecher, W. J. 1950. Convergent evolution in the American orioles. Wilson Bull., 62:51-86. 1951. Adaptations for food-getting in the American blackbirds. Auk, 68:411-440. BOUBIER, M. 1925. L'evolution de I'ornithologie. ii + 307 pp. Paris: Felix Alcan. Brehm, a. E. 1861. Das Leben der Vogel. xx + 707 pp. Glogau. 1864-1869. Illustrirtes Thierleben. 6 vols. Hildburghausen: Bibl. Inst. Bruch, C. F. 1828. Ornithologische Beitriige. Isis von Oken, 21:718-733. 1843. (Section fiir Zoologie, vierte Sitzung, am 23 September 1842) Deutsch. Naturf. Versamml. Ber., 1843:209-210. BULLER, W. L. 1872-1873. A History of the Birds of New Zealand, xxiv + 384 pp. London. Burt, W. H. 1930. Adaptive modifications in the woodpeckers. Univ. Calif. Publ. Zool., 32:455-524. Chapix, J. P. 1932-1939. The Birds of the Belgian Congo. 2 vols. Bull. Amer. Mus. Nat. Hist., Nos. 65 and 75. Cole, L. J. 1902. Suggestions for a method of studying the migrations of birds. Rept. Michigan Acad. Sci., 1902:67-70. Cooke, W. W. 1888. Report on the Bird Migration in the Mississippi Valley in the Years 1884 and 1885. Bull. U. S. Bur. Biol. Surv., No. 2, 313 pp. Washington, D. C. Cope, E. D. 1871. Synopsis of the extinct batrachia, reptilia and aves of North America. Trans. Amer. Philos. Soc, n.s., 14:1-252 + viii. Aves section, pp. 236-240. 1876. On a gigantic bird from the Eocene of New Mexico. Proc. Acad. Nat. Sci. Phila., 28:10-11. Cory, C. B., C. E. Hellmayr, and H. B. Conover 1918-1949. Catalogue of Birds of the Americas. Publ. Field Mus. Nat. Hist., Vol. 13, Pts. 1-11. (Publ. in 15 vols.) COUES, E. 1872. Key to North American Birds, viii + 361 pp. Salem. Dames, W. 1884. Ueber Archeoptet-yx. Palaeont. Abhandl., 2:117-196. 552 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Darwin, C. 1859. On the origin of species by means of natural selection, x + 502 pp. London: J. Murray. Delacoue, J. 1951. The Pheasants of the World. 347 pp. London: Country Life Ltd. Dickey, D. R., and A. J. van Rossem 1938. The Birds of El Salvador. Field Mus. Nat. Hist. (Zool. Ser.), 23:1-635. DOIJZHANSICY, T. 1937. Genetics and the Origin of Species, xviii + 446 pp. New York: Columbia Univ. Press. Dresser, H. E. 1871-1890. A history of the birds of Europe, including all species inhabiting the west- ern palaearctic region. 9 volumes. London. Drost, R. 1950. Study of bird migration 1938-1950. Proc. 10th Interuat. Ornithol. Congr., 216- 240. Elliot, D. G. 1864-1865. A Monograph of the Tetraoninae: or, Family of the Grouse, xx + 52 pp., 27 col. pis. New York. 1870-1872. A Monograph of the Phasianidae or Family of the Pheasants. 2 vols. New York. 1877-1882. A Monograph of the Bucerotidae, or Family of the Hornbills. xxxii + (2) + 146 pp., 60 pis. (57 col.). London. 1895. North American Shore Birds, a History of the Snipes, Sandpipers, Plovers and Their Allies. . . . xvi + 17-268, 72 pis. New York. Engels, W. L. 1940. Structural adaptations in thrashers (Mimidae: Genus Toxostoma) with com- ments on interspecific relationships. Univ. Calif. Publ. Zool., 42:341-400. Earner, D. S. 1950. The annual stimulus for migration. Condor, 52:104-122. FiNSCH, 0. 1867-1868. Die Papagaien. 2 volumes. Leiden. PiNSCH, O., and G. Hartlaub 1870. Die Vogel Ost-Afrikas. x + 897 pp. Leipzig. Fisher, H. I. 1946. Adaptations and comparative anatomy of the locomotor apparatus of New World vultures. Amer. Midi. Nat, 35:545-727. Friedmann, H. 1929. The Cowbirds. xvii + 421 pp. Springfield, 111. and Baltimore, Md. : C. C. Thomas. 1948. The Parasitic Cuckoos of Africa. Wash. Acad. Sci. Monogr. No. 1. 204 pp. FiJRllRINGER, M. 1888. Untersuchungen zur Morphologie und Systematik der Vogel, zugleich ein Beitrag zur Anatomie der Stiitz-und Bewegungsorgane. 2 vols, xlix + 1751 pp. Amsterdam: van Holkema. Gadow, H. F. 1891. Vogel. In Bronn's Klassen und Ordnungen des Thier-Reichs, Vol. 6. 1008 pp. Leipzig: C. F. Winter. SIBLEY: ORNITHOLOGY 653 Garroi), a. H. 1873a. On the carotid arteries of birds. Proc. Zool. Soc. London, 1873:147-472. 1873b. On certain muscles of the thigh in birds and their value in classification. Pt. I. Proc. Zool. Soc. London, 1873:626-644. 1874. On certain muscles of the thigh in birds and their value in classification. Pt. II. Proc. Zool. Soc. London, 1874:111-123. Gatke, H. 1891. Die Vogelwarte Helgoland, xii + 609 pp. Braunschweig. Glenny, F. H. 1940. A systematic study of the main arteries in the region of the heart. Aves. Anat. Rec, 76:371-380. Gloger, B. von 1833. Das Abandern der Vogel durch Einfluss des Klima's. xxix + 159 pp. Breslau: Schulz and Co. Gould, J. 1840-1848. The Birds of Australia. 7 vols., 600 col. pis. London. 1851-1869. (Suppl. to above.) iv + 158 pp., 81 col. pis. London. 1849-1861. A Monograph of the Trochilidae, or Family of Humming-Birds. 5 vols. London. Grasse, p. p. et al. 1950. Oiseaux. Vol. 15 of Traite de Zoologie, Anatomie, Systematique, Biologic. 1164 pp. Paris: Masson et cie. Groebbels, F. 1932-1937. Der Vogel. 2 vols. Berlin: Gebr. Borntraeger. Harris, H. 1928. Robert Ridgway. Condor, 30:5-118. Hartert, E. 1903-1923. Die Vogel der palaarktischen Fauna. 3 vols. Berlin and London. Hartert, E., H. F. Witherby, F. C. R. Jourdain, and N. F. Ticehurst 1912. A Hand-List of British Birds, with an Account of the Distribution of Each Species in the British Isles and Abroad, xii + 237 pp. London. Hartlaub, G. 1857. System der Ornithologie Westafrica's. Ixvi + 280 pp. Bremen. Heilmann, G. 1926. The Origin of Birds, vi + 208 pp. London: Witherby. Heinroth, O. 1910. Beitrage zur Biologic, namentlich Ethologie und Psychologie der Anatiden. Verb. 5te. Ornithol. Kongr., pp. 589-702. Heinroth, O., and M. Heinroth 1926-1928. Die Vogel Mitteleuropas. 3 vols. Berlin: Hugo Bermiihler. 1941. Das Heimfinde-Vermogen der Brieftaube. Journ. Ornithol., 89:213-256. Hellmayr, C. E. 1918-1949. [See Cory, C. B., C. E. Hellmayr, and H. B. Conover.] Herrick, F. H. 1901. The Home Life of Wild Birds, xix + 148 pp. New York and London: Knicker- bocker Press. 654 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES HOFER, H. 1949. Die Gaumenliicken der Vogel. Acta Zool. Internat. Tidskr. Zool., 30:209-248. HOMEYER, E. F. VON 1881. Die Wanderungen der Vogel mit Rlicksicht auf der Ziige der Saugethiere, Fische und Insecten. 415 pp. Leipzig. 1885. Verzeichniss der Vogel Deutschlands. 16 pp. Vienna. Howard, H. E. 1907-1915. The British Warblers. 2 vols, in 10 pts. London: R. H. Porter. 1920. Territory in Bird Life, vi + 308 pp. London: John Murray. Howard, Hildegarde 1947. A preliminary survey of trends in avian evolution from Pleistocene to Recent time. Condor, 49:10-13. 1950. Fossil evidence of avian evolution. Ibis, 92:1-21. Hudson, G. E. 1937. Studies on the muscles of the pelvic appendage in birds. Amer. Midi. Nat., 18:1-108. 1948. Studies on the muscles of the pelvic appendage in birds. 2: The heterogeneous order Falconiformes. Amer. Midi. Nat., 39:102-127. Hudson, W. H. 1920. Birds of La Plata. 2 vols. London. H€:xLEY, T. H. 1867. On the classification of birds ; and on the taxonomic value of the modifications of certain of the cranial bones observable in that class. Proc. Zool. Soc. London, 1867:415-472. Ising, G. 1945. Die physikalische Moglichkeit eines tierischen Orientierungssinnes auf Basis der Erdrotation. Arckiv f. Matematik, Astron. och Fysik, Bd. 32A. No. 18. 23 pp. Kessler, Karl F. 1853. Einige Beitrage zur Wanderunggeschicte der Zugvogel. Bull. Soc. Nat. Moscou, 26(1): 166-204. Kleinschmidt, 0. 1900. Arten oder Formenkreise. Journ. f. Ornithol., 48:134-139. 1926. Die Formenkreislehre und das Weltwerden des Lebens. x + 188 pp. Halle. 1930. Ibid. Trans, into English as: The Formenkreis Theory and the Progress of the Organic World. London: H. F. and G. Witherby. Knowlton. F. H. 1909. Birds of the World, xiii + 873 pp. New York. Kramer, G. 1949. tJber Richtungstendenzen bei der nachtlichen Zugenruhe gekafigter Vogel. Ornithol. als Biol. Wiss., pp. 269-283. Heidelberg: Carl Winter. 1950. Eine neue Methode zur Erforschung der Zugorientierung und die bisher damit erzielten Ergenbnisse. Proc. 10th Internat. Ornithol. Congr., pp. 269-280. Lack, D. 1939. The behaviour of the robin. Pts. I and II. Proc. Zool. Soc. London, 109 A: 169- 219. SIBLEY: ORNITHOLOGY 655 Lambrecht, K. 1933. Handbuch der Palaeornithologie. xix + 1024 pp. Berlin: Gebr. Borntraeger. LORENZ, K. 1931. Beitrage zur Ethologie sozialer Corviden. Journ. f. Ornithol., 79:67-120. 1935. Der Kumpan in der Umwelt des Vogels. Journ. f. Ornitliol., 83:137-213, 289-413. 1937. The companion in the bird's world. Auk, -54:245-273. Lowe, P. R. 1944. An analysis of the characters of Archaeopteryx and Archaeornis. Were they reptiles or birds? Ibis, 86:517-543 (figs. 5-10). Marsh, 0. C. 1870. Notice of some fossil birds, from the Cretaceous and Tertiary formations of the United States. Amer. Journ. Sci., ser. 2, 49:205-217. 1872. Preliminary description of Hesperornis regalis, with notices of four other new species of Cretaceous birds. Amer. Journ. Sci., ser. 3, 3:360-365. 1880. Odontornithes: A monograph on the extinct toothed birds of North America. Rept. Geol. Expl. 40th Parallel, pp. 1-201. Mathews, G. M. 1910-1928. The Birds of Australia. 12 vols. London. Mayr, E. , 1935. Bernard Altum and the Ten-itory theory. Proc. Linn. Soc. New York, 45-46: 24-38. 1942. Systematics and the Origin of Species, xiv + 334 pp. New York: Columbia Univ. Press. 1950. Speciation in birds. Progress report on the years 1938-1950. Proc. 10th Internat. Ornithol. Congr., pp. 91-131. Mayr, E., and D. Amadon 1951. A Classification of Recent Birds. Amer. Mus. Nov., no. 1496, 42 pp. Mayr, E., E. G. Linsley, and R. L. Usinger 1953. Methods and Principles of Systematic Zoology, vii + 328 pp. New York: McGraw-Hill. McDowell, S. 1948. The bony palate of birds. Pt. I. The Palaeognathae. Auk, 65:520-549. Mendel, G. 1865. Versuche iiber Pflanzen-Hybriden. Verb. Naturf. Ver. Briinn. Bd. 4. Miller, A. H. 1937. Structural modifications in the Hawaiian goose (Nesochen sandvicensis) a study in adaptive evolution. Univ. Calif. Publ. Zool., 42:1-80. 1941. Speciation in the avian genus Junco. Univ. Calif. Publ. Zool., 44:173-434. Miller, L. H. 1909. Pavo californicus, a fossil peacock from the Quaternary asphalt beds of Rancho La Brea. Univ. Calif. Publ. Dept. Geol., 5:439-448. Miller, L. H., and Ida DeMay 1942. The fossil birds of California. Univ. Calif. Publ. Zool., 47:47-142. Milne-Edwards, A. 1867-1871. Recherches anatomiques et paleontologiques pour servir a I'histoire des oiseaux fossiles de la France. Vol. I, 474 pp.; vol. II, 632 pp. Atlas, 200 pis. in 2 vols. Paris: Masson et cie. 656 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Morgan, C. Lloyd 1896. Habit and Instinct. 351 pp. New York and London: Edward Arnold. MULLER, J. P. 1847. Nachtrag zu der Abhandlung liber die Stimmorgane der Passerinen. Arch. Anat. Physiol., 1847:397-399. Murphy, R. C. 1936, Oceanic Birds of South America. 2 vols. Amer. Mus. Nat. Hist. New York. Newton, A. 1893-1896. A Dictionary of Birds. 1088 pp. London. Newton, E. T. 1886. On the remains of a gigantic species of bird {Gastornis klaasseni, n. sp.) from the Lower Eocene beds near Croydon. Trans. Zool. Soc, 12:143-160. Nice, Margaret M. 1937. Studies in the life history of the song sparrow. I. Trans. Linn. Soc. New York, 4:1-247. 1943. Studies in the life history of the song sparrow. II. Trans. Linn. Soc. New York, 6:1-328. 1941. The role of territory in bird life. Amer. Midi. Nat., 26:441-487. Niethammer, G. 1937-1942. Handbuch der Deutschen Vogelkunde. 3 vols. Leipzig: Akademische Verlagsgesellschaft M.B.H. Nopcsa, F. 1907. Ideas on the origin of flight. Proc. Zool. Soc. London, 1907:223-236. Oates, E. W., and W. T. Blanford 1889-1898. The Fauna of British India, including Ceylon and Burma. Birds. 4 vols. London. Odum, H. T. 1948. The bird navigation controversy. Auk, 65:584-597. Oliver, W. R. B. 1930. New Zealand birds. Wellington, N. Z. Owen, R. 1863. On the Archeojiteryx of von Meyer, with a description of the fossil remains of the longtailed species from the lithographic stone of Solnhofen. Phil. Trans- act. London, 33-47. 1866-1868. On the Anatomy of Vertebrates. Vol. 2. viii + 592 pp. London: Longmans, Green and Co. Palmen, J. A. 1876. Ueber die Zugstrassen der Vogel. 292 pp. 8 vols. Leipzig: W. Engelmann. Peters, J. L. 1931-1951. Check-list of Birds of the World. 7 vols. Cambridge, Mass.: Harvard Univ. Press. Petronievics, B. 1921. Ueber das Becken, den Schultergiirtel und einige andere Telle der Londoner Archaeopteryx. Genf., 1-31. Phillips, J. C. 1922-1926. A Natural History of the Ducks. 4 vols. Boston. SIBLEY: ORNITHOLOGY 657 PyCEAFT, W. p. 1910. A History of Birds. 31 + 458 pp. London. Reichenow, a. 1875. In "Protokoll der constituirenden Friijahrs-Versammlung." Journ. f. Ornithol., 23:347. 1893. System und Genealogie. Ornithol. Monatsber., 1:113-117. Rensch, B. 1929. Das Prinzip geographisclier Rassenkreise und das Problem der Artbildung. Berlin: Gebr. Borntraeger. Richardson, F. 1942. Adaptive modifications for tree-trunk foraging in birds. Univ. Calif. Publ. Zool., 46:317-368. RiDGWAY, R. 1869. Notices of certain obscurely known species of American birds. Proc. Acad. Nat. Sci. Phila., 21:125-135. 1881. Nomenclature of North American birds chiefly contained in the United States National Museum. Bull. U. S. Nat. Mus., 21:1-94. RiDGWAY, R., and H. Friedmann 1901-1950. The Birds of North and Middle America. Bull. U. S. Nat. Mus., Vol. 50, pts. 1-11. Rothschild, Miriam, and Teresa Clay 1952. Fleas, Flukes and Cuckoos, xiv + 304 pp. New York: Collins. Rowan, W. 1931. The Riddle of Migration, xiv + 151 pp. Baltimore: Williams and Wilkins. Rydzewski, W. 1950. Bird-Ringing Schemes Known to Be Operating at Present. Proc. Tenth Internat. Ornithol. Congr., pp. 356-359. Salvin, O., and F. D. Godman 1879-1904. Biologia Centrali-Americana. Aves. 4 vols. London. Saunders, H. 1889. An Illustrated Manual of British Birds. 40 -f 754 pp. (1st ed.) London. 1899. An Illustrated Manual of British Birds. 40 + 776 pp. (rev. ed.) London. Schlegel, H. 1828. Verhandeling ter Beantwoording der Vrage: "Daar er nog veel Duisterheid en Verschil van Gevoelens Platts heeft omtrent de Gewesten, waarheen zich de bij ons bekend gewordene Trekvogels begeven, etc." Vern. holl. Maat. Wet. Haarlem (2), 16:129-292. 1844a. (See Siebold, 1844.) 1844b. Kritische Uebersicht der Europaischen Vogel. 2 pts., pt. 1, 135 pp; pt. 2, 116 pp. Leiden and Paris. ScLATER, P. L., and W. H. Hudson 1888-1889. Argentine Ornithology. A Descriptive Catalogue of the Birds of the Argentine Republic. 2 vols. London. Seebohm, H. 1887. The Geographical Distribution of the Family Charadriidae, or the Plovers, Sandpipers, Snipes, and Their Allies, xxix + 524 pp. London. 658 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Selous, E. 1901. Bird Watching, xi + 347 pp. London. 1905a. Bird Life Glimpses, viii + 355 pp. 1905b. The Bird Watclier in the Shetlands. x + 388 pp. Sharpe, R. B. 1899-1909. Hand-List of the Genera and Species of Birds in the British Museum. 5 vols. London. Sharpf;, R. B., and others 1874-1898. Catalogue of the Birds in the British Museum. 27 vols. London. Shelley, G. E. (and, in part) W. L. Sclater 1896-1912. The Birds of Africa, Comprising All Species Which Occur in the Ethiopian Region. 5 vols. 502 pp. London. Shufeldt, R. W. 1891. Fossil birds from the Equus beds of Oregon. Amer. Nat., 25:818-821. Siebold, Pii. Fr. de 1844-1850. Fauna Japonica: Aves: Temminck, C. J., and Schlegel, H. Lugduni Batavorum, Apud A. Arnz et socios. Steiner, H. 1917. Das Problem der Diastataxie des Vogelfltigels. Jenaische Zeitschr. f. Naturwiss., 55:221-496. Stolpe, M. 1932. Physiologisch-anatomische Untersuchungen iiber die hintere Extremitat der Vogel. Journ. f. Ornithol., 80:161-247. Stresemann, E. 1927-1934. Aves-Vogel. In Kiikenthal und Krumbach, Handbuch der Zoologie, Bd. 7 (2te Halfte). 899 pp., 944 figs. Berlin: De Gruyter. 1951. Die Entwicklung der Ornithologie. xv + 431 pp. Berlin: F. W. Peters. Strong, R. M. 1939-1946. A Bibliography of Birds. Field Mus. Nat. Hist. (Zool. Ser.), 25, pts. 1-3. SwANN, H. K., and A. Wetmoke 1924-1945. A Monograph of the Birds of Prey. London. SWINTOX, W. E. 1934. A Guide to the Fossil Birds, Reptiles and Amphibians in the Dept. Geol. and Palaeont. Brit. Mus. (Nat. Hist.). London. Thomson, J. A. 1923. The Biology of Birds, xi + 436 pp., 9 pis., 59 figs. London. TlNMlCRGEN, N. 1936. The problem of sexual fighting in birds; and the problem of the origin of "territory." Bird Banding, 7:1-8. 1939a. The behavior of the Snow Bunting in spring. Trans. Linn. Soc. New York, 5:1-95. 1939b. On the analysis of social organization among vertebrates, with special refer- ence to birds. Amer. Midi. Nat., 21:210-234. 1942. An objective study of the innate behaviour of animals. Bibl. Biotheor., 1:39-98. 1948. Social releasers and the experimental method required for their study. Wilson Bull., 60:6-52. 1951. The Study of Instinct, xii + 228 pp. London: Oxford Univ. Press. SIBLEY: ORNITHOLOGY 659 TiNBEBGEN, N., and A. C. Perdeck 1950. On the stimulus situation releasing the begging response in the newly hatched Herring Gull chick {Larus a. argentatus Pont.)- Behaviour, 3:1-38. Watson, J. B., and K. F. Lashley 1915. An historical and experimental study of homing. Pap. Dept. Mar. Biol. Carnegie Inst., 7:9-83, 6 pis., 9 figs. Wetmobe, a. 1921. A fossil owl from the Bridger Eocene. Proc. Acad. Nat. Sci. Phila., 73:455-458. 1940. A check-list of the fossil birds of North America. Smithsonian Misc. Coll., 99 (no. 4): 1-81. 1950. Presidential Address. Recent Additions to Our Knowledge of Prehistoric Birds 1933-1949. Proc. 10th Internat. Ornithol. Congr., 51-74. White, G. 1789. The natural history of Selborne. v + 468 + (13). London. Withebby, H. F., F. C. R. Jourdain, N. F. Ticehurst, and B. W. Tucker 1943. The Handbook of British Birds. (Rev. ed.). 5 vols. London: H. F. and G. Witherby. WOLFSON, A. 1945. The role of the pituitary, fat deposition, and body weight in bird migration. Condor, 47:95-127. Wood, C. 1931. An Introduction to the Literature of Vertebrate Zoology, xix + 643 pp. London: Oxford Univ. Press. Yarrell, W. 1837-1843. A History of British Birds. 3 vols. London. Yeagley, H. L. 1947. A preliminary study of a physical basis of bird navigation. Journ. Applied Physics, 18:1035-1063. ZlMMEB, J. T. 1931. Studies of Peruvian Birds. I. Amer. Mus. Nov., no. 500, pp. 1-23. MAMMALOGY IN NORTH AMERICA By W. J. HAMILTON, JR. Cornell University From the dawn of history, mammals have played a vital part in the destiny of man. The mammal fauna of North America has been of tremendous economic significance, in one manner or another, to human populations. As food for the earlier settlers, many species provided, and continue to do so, a source of meat of not inconsiderable quantity. The peltries of our fur-bearers supply a substantial revenue to the trapper. Once the primary fur animal of the continent, the beaver influenced the exploration and settlement of the West and the northern latitudes. Esthetic values are not so tangible, but are evident in the hordes of tourists who annually visit our national parks to see the great bears and hoofed species as well as the attractions of the geysers, waterfalls, and other natural phenomena. On the other hand, the losses sustained through destruction of crops and foodstuffs by mammals may be very great. Some species play a major role in the trans- mission of disease organisms, such as sylvatic plague, murine typhus, spotted fever, rabies, and others of lesser importance. These economic relations have inspired extensive studies, through which much has been learned regarding the habits of certain species. The results of these investigations are continually being catalogued. Research in the field has not kept pace with that accomplished on some of the other classes of animals, for mammals are often shy and retiring in their habits and many are nocturnal, making observation difficult. The study of mammals needs no economic justification, although pure research has been repeatedly applied to factors which relate to man's welfare. This has been aptly expressed by Miller (1928). There is nothing to be gained by denying that discovery for its own sake has always been the mainspring of work in all branches of scientific endeavor, including mammalogy. . . . This incentive requires no other apology than an indication of how the knowledge thus gained has contributed to human advancement. Indeed, an understanding of the relationships between the obscure seeker after facts and man's well-being must forever justify the worker in pure research. The science of mammalogy may be said to date back only to the time of Lin- naeus. Prior to the middle of the eighteenth century, the study of these animals had lacked conciseness. The binomial system of Linnaeus, however simple it may appear to present-day students, proved so useful a tool that it seems impossible that any serious study of animals or plants could have proceeded without it. In his tenth edition of the Systema Naturae, published in 1758, Linnaeus in- cluded only 86 mammals. A century later, 220 kinds were known to North America alone. Presently, nearly 3,000 species and subspecies are recognized as occurring north of Panama. North of the Mexican boundary, nearly 400 full species are recognized today, some of these containing 40 subspecies alone. The number of [661] 662 A CENTURY Of PROGRESS IN THE NATURAL SCIENCES JOSEPH LEIDY 1823-1891 SPENCER FULLERTON BAIRD 1823-1887 CLINTON HART MERRIAM 1855-1942 HENRY FAIRFIELD OSBORN 1857-1935 HAMILTON: MAMMALOGY IN NORTH AMERICA 663 fossil mammals that have been described almost equals that of living forms. It is presumed that when all the races are described, more than 20,000 will have been recorded from the entire world. Few reference works were available to the early American biologists who had an interest in mammals. Richard Harlan, a close friend and supporter of Audubon, in 1825, published the first installment of his Fauna Americana, which treated mammals exclusively. While it was principally a compilation, based in large measure on Desmarest's Mammalogie, it served a useful purpose for the time. The following year John B. Godman's North American Natural History, or Mastology, lent further impetus to the study of mammals. There is much on the habits of the commoner species in tliis report. The first part of DeKay's Zoology of New York, dealing with the mammals, was published in 1842. This work in- cludes considerable discussion of extra-limital species, and is a useful historical account. For the first substantial report on the mammals of North America, we are indebted to Audubon and Bachman. The Vivi2)arous Quadrupeds of NoHh Amer- ica appeared from 1846 to 1854. The plates, with a few exceptions, had been previously published in large oblong folio, without text, commencing as far back as 1840. The three volumes included 197 species, exclusive of varieties, of which about 160 were figured. John Bachman has seldom been properly credited for his great contribution to American mammalogy. A lifetime spent in the ministry, he yet found time to make lasting contributions to science. His friendship with Audubon dated from 1831 until the latter's death twenty years later. Dr. Bachman was a learned zoologist of his day. In 1839, Audubon and he began work on the great Quadru- peds. Audubon was never to see the completed work, dying in 1851 when the first volume had been completed. His sons, John and Victor, were to color the plates and arrange for the editing and sales, but the greatest share would fall to Bachman, who was to make the dissections, write the systematic accounts and contribute largely to the text, through his vast knowledge of the life histories of the commoner species. Bachman had a restraining influence on his friend, cau- tioning Audubon repeatedly to exercise care in his spontaneity. In 1840, Bach- man addressed his friend Audubon thus : When we meet, we shall talk about the partnership in the quadrupeds. I am willing to have my name stand with yours, if it will help the sale of the book. The expenses and the profits shall be yours or the boys. I am anxious to do something for the benefit of John and Victor, in addition to the treasures I have given them [Bachman was the father- in-law of Audubon's sons]. . . . Don't flatter yourself that the quadrupeds will be child's play. I have studied them all my life. We have much, both in Europe and America, to learn on this subject. The skulls and the teeth must be studied, and the color is as variable as the wind; down, down in the earth they grovel, while we, in digging and studying, may grow old and cross. Our work must be thorough. I would as soon stick my name to a forged Bank Note as to a mess of Sloupviaigre. Present-day students of mammalian life histories critically examine, or should do so, the pages of the Quadrupeds before commencing a serious study of any species. The difficulties of vertebrate research in the early nineteenth century, par- ticularly the review of literature and access to museum specimens, are set forth in the introduction of the Quadrupeds. The young field naturalist will profit from reading this account. 564 a century of progress in the natural sciences Baird and the Smithsonian At its inception, the Smithsonian Institution was charged with the responsi- bilit}^ for maintaining a museum. Spencer Fullerton Baird, then assistant to Secretary Henry submitted a report, detailing the need of research and publica- tions that would accrue from such investigations. This was in accord with Henry's view. The genius of Baird and his inspiration to the young collectors under him has not been fully appreciated. Baird had an enthusiasm and matchless knowledge of the vertebrates that will seldom be equaled. In 1853, Congress had appropri- ated $150,000 to defray the expenses of the survey of the various routes along which it was supposed that a railroad might be constructed from the Mississippi River to the Pacific. For this purpose, six parties were organized by the War Department. Through the efforts of Baird, persons capable of making collections and observations in natural history were assigned to these parties. These expedi- tions resulted in the most voluminous collections of the time. Earlier Wilkes (1838-1842) and his associates had made collections on the U. S. Exploring Expe- dition. Baird's study of these collections, particularly the mammals, was precise and stands as a monument to his untiring industry (Baird, 1857). While Baird presumably cared for the mammal collection until 1879, the U. S. National Museum was organized in that year by G. Brown Goode, under the instruction and guid- ance of Baird. Dr. Elliott Coues, distinguished ornithologist and mammalogist, was designated as curator of mammals. His Fur-Bearing Animals, a monograph of the North American Mustelidae published in 1877, was a classic of the time and is of lasting value. Frederick W. True, renowned for his studies on cetaceans, was curator of mammals from 1881 to 1908. Gerrit S. Miller, Jr., is indelibly stamped in the minds of mammalogists for his North American Recent Mammals (Miller, 1924), the only check list of North American mammals presently available to the student. In this report synonomy, type locality, and distribution are given. While now an outdated reference work, it is still of considerable value to the student. Remington Kellogg became curator of mammals upon Miller's retirement. His knowledge of vertebrates is unsurpassed. He has published in many fields, but his greatest contributions have been on cetaceans. Kellogg's place as a master zoologist was recognized in 1948 when he was made director of the U. S. National Museum. For a fuller account of the Smithsonian, the reader is referred to Kellogg (1946). The Influence of Merriam on American Mammalogy Clinton Hart Merriam had a profound effect upon mammalogy, indeed he was preeminent in the field. His accomplishments and influence on others will long be felt in American zoology. As a youngster in upstate New York, his passion for birds and mammals resulted in substantial early reports. Upon the comple- tion of his medical school studies in 1879, Dr. Merriam practiced for six years in Locust Grove, but his growing interest in mammals was evident during this period. In 1884 his Mammah of the Adirondaks was published. This report set a new standard, embodying for the first time details of life histories that have seldom been surpassed in a local work. A year earlier he had begun correspond- HAMILTON: MAMMALOGY IN NORTH AMERICA 665 ence with young Vernon Bailey, a Minnesota farm boy. This lad, later to become Merriam's brother-in-law, was an indefatigable collector. Through Bailey's well prepared specimens and large series of the less common species (at least in col- lections), Merriam may have been first encouraged to consider the possibility of a country-wide survey of mammals. As with so many naturalists, Merriam's first love was ornithology. The found- ing of the American Ornithologists' Union in 1883 Ijrought him in contact with the masters of the day, including Baird, Bendire, J. A. Allen, Ridgway, and others. He was elected secretary of the society. In 1885, through the efforts of the A.O.U., Congress authorized the establishment of a section of ornithology to be a branch of the Division of Entomology, then under the Commissioner of Agriculture. Merriam was appointed as ornithologist in this newly created section. He gave up the practice of medicine and assumed the duties that were to play so important a part in North American mammalogy. His fellow student in medical school. Dr. A. K. Fisher, was invited as assistant ornithologist. Most of us remember Fisher best for his Hawks and Owls of the United States, published in 1893. Within three years, the section became the Division of Economic Ornithology and Mammalogy. In 1905, the Bureau of Biological Survey was founded, an outgrowth of the smal- ler unit. We now know this bureau as the Fish and Wildlife Service, under the Department of the Interior. Merriam assembled a group of able men for the Bureau, and sent collectors into the unexplored West. He inaugurated the technical North American Fauna series, revisions and description of mammals occupying many of these important publications. By the early 'nineties, Merriam had planned his life work; studies that would determine some of the factors which limit the distribution of plants, birds, and mammals. His descriptions of new mammals, including several dis- tinctive genera, may be partially credited to the industry of Bailey, who was sending to Washington scores of undescribed forms. The San Francisco Mountains of Arizona offered a splendid opportunity to study altitudinal distribution. The report of this trip gave a clue to his later reports on distribution (Merriam and Stejneger, 1890). However modified in later years, the Arizona study was fundamental. Many may disagree with his temperature laws, but in parts of North America these have stood the test of time. To be sure, there are valid objections to these ''temperature summations," but they appear to hold in a great part of western North America. The standards of Merriam were of the highest caliber. However harsh he might appear to some, he gave freely of advice and aided many an aspiring youngster. Recently I have seen his entire correspondence to one of his field assistants, a collector of no mean ability. When this assistant offered to resign, feeling that he had been accused of misusing government property, Merriam wrote in longhand, on plain paper, the following letter, dated June 14, 1894. Don't lose your head, even if the provocation seems great — from your standpoint. It is evident that I was mistaken as to what you actually did. I thought you had sawed up or made a packing box of the two trays from the new chest we sent you last — not dreaming that you had kept two trays of the old chest with you so long. Please bear in mind that I am held personally responsible to the Department for all property belonging to the Division, and am now charged with several hundred dollars worth of property that has gone in the field and not likely to be returned. The most important single thing for a young man to learn is self control — without ()(,(i A CENTURY OF PROGRESS IN THE NATURAL SCIENCES this he cannot hope to fill a useful field among his fellow men. If you ever get so very mad you feel you must write an impudent letter, the best way is to sit right down and write it and say all the mean things you can think of. Then take the letter and your hat, having relieved your mind, and take a walk to some secluded spot. Then take out your match box and set fire to the letter and stay by it until it has been decomposed into its chemical constituents. Whatever you do, don't ever mail such a letter — particularly in an oflicial capacity. Furthermore, don't mix personal and official matters in the same letter. Always write as freely as you wish about personal things, only not on the same sheet with your official letters which go on file.^ — C. H. M. The influence of Merriam on j^ounger naturalists of the time cannot be denied. His greatest student was Vernon Bailey, a heroic figure in American mammalogy. Many "unknowns," later to become celebrated for their own researches, collected for him. E. W. Nelson, E. A. Goldman, and W. H. Osgood may be numbered among his illustrious "students." Dr. Nelson later served as chief of the Biological Survey (1916-1927), Goldman is noted for his Mexican surveys, and Wilfred H. Osgood was director of zoology at the Chicago Natural History Museum at the time of his death. For a detailed account of Merriam,^ the reader should see the stimulating account by Osgood (1943). Nearly 500 publications, many of mono- graphic scope, are listed by Grinnell (1943). The United States Biological Survey No other organization has played such an outstanding role in American wild- life as has the U. S. Biological Survey. Its function is the investigation of life histories, habitats, ranges, distribution, and the economic, recreational, cultural, and other values of American birds and mammals. Over the years, a major em- phasis has been placed on the repression of noxious rodents and predatory mam- mals where such was needed. The vast number of scientific publications detailing the researches conducted by this agency is without parallel. When the first appropriation for a Branch of Economic Ornithology in the Division of Entomology was made in 1885, Americans were at long last becoming conscious of the increasing plight of our wildlife resources. They had seen the fate of the buffalo determined with completion of the Union Pacific. Ribbons of steel had separated the great beasts into a northern and southern herd, and the railroad provided the needed transport for the spoils of the hide hunters. Unwise introductions of exotics and the scandalous slaughter of wildlife had the effect of focusing attention on the plight of this great natural heritage. In its second year, the division took cognizance of mammals, primarily in their relation to agriculture and horticulture. It appears that Dr. Merriam had little use for the term "economic," and his leadership led to a steady subordination of the practical problems to those of the scientific. It was not long before his interests prevailed. Studies in geographic distribution, which Merriam considered equally or more important than the economic, took precedence over the practical. Economic and agricultural publications were to be published in the form of special reports or circulars (the familiar Farmers' Bulletin), while the scientific was to be brought out in the North American Fauna series. From 1891 until 1906, geographic dis- tribution was the keynote of research, with economic relations playing a lesser role. This trend was reflected in the Secretary of Agriculture's report for 1890, in which he declared •, HAMILTON: MAMMALOGY IN NORTH AMERICA 667 The name of this Division is unfortunate as it conveys an erroneous idea of the nature of its work. The division is in effect a biological survey, and should be so named, for its principal occupation is the preparation of large-scale maps of North America, showing the boundaries of the different faunas and floras, or life areas. The results of these explorations bore fruit in 1894, when the divisional report for that year announced that the problem of temperature control of the geo- graphic distribution of animals and plants had been solved. The Weather Bureau had provided temperature data which, when plotted on the biogeographic maps, conformed with a high degree of exactness to the boundaries of the life zones as established by Merriam. For a decade, the biological exploration of North America continued. The geographic distribution of species in the West received major attention until 1906, when the Bureau of Biological Survey, as it was now called, again shifted its emphasis to economic problems. Merriam had selected his staff with care. His counsel and training of the young field agents did not go unrewarded. To one of his younger field naturalists, J. Alden Loring, Merriam wrote more than a score of letters in a matter of eight months. These are replete with instructions, criti- cism of skins, and helpful advice. It is presumed he carried on as lively a corre- spondence with his other field assistants. When he was not in the field, Merriam found time to initiate the Fauna series. From 1889 to 1896, this indefatigable scientist authored the first eleven of the fanual series, all of monographic scope. These were the first revisions and serious taxonomic studies ever made on North American mammals. They stand as a monument to ]\Ierriam's industry and taxo- nomic judgment. The advance of mammalogy at this time was fortunately not dependent on the resources of the government. In 1899, Edward H. Harriman organized and financed an expedition to Alaska, members of the Biological Survey sharing in the investigation. In succeeding years, the scope was enlarged to include Canada and Mexico. In 1907, Congressional hearings resulted in partial abandonment of the dis- tributional studies. More emphasis was expended on practical pursuits. The well known reports of Professor David Fj. Lantz now appear. Many of these are con- cerned with injurious rodents and measures for their control. The undercurrent of public opinion that dictated this shift to a practical point of view was a sound one. With the amazing growth of agriculture and the conse- quent increase in the value of its products, information was sorely needed on the control of the many pests which took a huge annual toll. The agriculturist was no longer content with reports detailing the habits, distribution, and characters of the pests which pilfered his crops or destroyed his livestock. A new supply of food was available to the wolves and coyotes, and the stockmen took the brunt of this toll. An investigation of the wolf in relation to stock raising was published by Bailey (1907). This was followed by a shorter article by the same author, in which emphasis was placed on den hunting, with the subsequent destruction of the litter. Following the recommendations outlined in these reports, an estimated 1,800 wolves and 23,000 coyotes were accounted for in a single year. Not until 1915, with increasing depredation from predatory animals, did Congress relieve the Forest Service of this effort. With a sizable appropriation to the Survey, Congress directly ordered the destruction of "wolves, coyotes and other animals injurious to agriculture and animal husbandry on the national forests and the 668 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES public domain," thus placing the responsibility directly on the Survey. This action may have been precipitated by a disastrous outbreak of rabies among wild animals in the West. In 1916 more than half of the appropriation for food habits research was expended in activities to control the wolf and coyote. Efforts to eradicate these animals have been continued. Where conditions are suitable, poison is by far the most economical and effi- cient known agent for the destruction of the coyote, other predatory mammals, and rodents, where they are abundant. The continued use of this method in eradicating noxious species brought many objections. Many useful species were unintentionally killed. Valuable fur-bearers have been destroyed in considerable numbers. Continued protests by those who favored a reduction in poisoning opera- tions and a modified policy of control by a government agency culminated in open discussion of the pros and cons of the method. A symposium on predatory animal control was held in New York City, May 21, 1930, at which scientists of the Bio- logical Survey defended the program, while those from universities, museums, and other organizations brought out the dangers attendant on the widespread use of poison. For details of these discussions, the reader is referred to the August, 1930, issue of the Journal of Mammalogy. The shift of emphasis from surveys and distributional studies to that of con- trol of noxious pests was inevitable. Pressure from agriculture and livestock interests had brought this to pass. AVithout the purely scientific studies of the Merriam era, however, the distribution of various small mammals of economic significance would not have been known. When the call came for control, imme- diate steps could be taken and widespread efforts made at reduction. This is essential in controlling many of our western ground squirrels, for piecemeal efforts result only in temporary relief. Other divisions of the survey have been occupied with mammal investigations. This is reflected in the scope of the reports that have been published in recent years. Until rather recently, the Division of Food Habits Research, while empha- sizing the economic status of birds, had made marked contributions to our knowl- edge of wild mammal dietary. Such studies are essential in determining, in part, economic relationships. Considerable effort has been directed to the investigations and life histories of rodents, by far the major share of such studies being focused on the Norway rat. This unmitigated pest has no redeeming quality. The loss it occasions yearly to our foodstuffs and as an agent in the spread of disease is all too well known. Research directed toward new raticides has played a not incon- siderable part in our increasing and successful war against this arch enemy of man. Research on fur-bearers, with special emphasis on problems of the fur fanner, has long been under the Division of Fur Resources. These investigations are con- cerned primarily with nutritional and disease studies. The Federal Aid in Wildlife Restoration program was inaugurated in 1938 under the Pittman-Robertson Act, which provides for the use, in behalf of wild- life, of income from the Federal excise tax on sporting arms and ammunition. In the thirteenth year of the program, closing on June 30, 1951, a sum of $17,846,423 was made available for tliis work. Federal allotment is matched by a 25 per cent contribution from the states to carry out approved projects. Many of the state conservation departments and the state colleges and universities have profited by HAMILTON: MAMMALOGY IN NORTH AMERICA 669 the funds thus made available for research. In a recent year, 184 individual proj- ects were under way in 44 states, Alaska, Hawaii, Puerto Eico, and the Virgin Islands, with emphasis on game and fur animals. In 1951, 33 states had research projects on deer, 22 were investigating fur-bearer problems, chiefly muskrat and beaver, while 11 were investigating rabbits and hares. Other mammals that have received attention are antelopes, squirrels, mountain sheep and goats, elk, moose, and bison. One of the most detailed state mammal surveys yet undertaken has been supported by Pittman-Robertson funds. This Pennsylvania project, under the direction of J. K. Doutt of the Carnegie Museum, has provided more details regarding the distribution and habits of the mammals inliabiting a single com- monwealth than any previous study. Under provisions of President F. D. Roosevelt's reorganization plan, made effective June 30, 1940, the Bureau of Fisheries and the Bureau of Biological Survey, in the Department of the Interior, with their respective functions, were consolidated into one agency, to be known as the Fish and Wildlife Service. Progress in Paleontological Research Few areas are so rich in fossil mammals as western North America. The suc- cessive assemblages of animals which once lived in this vast area have been faith- fully studied for the past ninety years. John Evans, assistant to Dr. David D. Owen, Dr. F. V. Ilayden of the U. S. Geological Survey, and others led important expeditions into this unexplored region. Collecting was not the prosaic occupa- tion of today. Pack horses and wagons carried out the rewards of these expeditions to the single transcontinental railroad; hostile Indians made these explorations extremely hazardous. Joseph Leidy was to lay the foundation for the science of American paleon- tology. Trained in medicine. Dr. Leidy had little time to devote to practice, the consuming interest in fossils occupying ever more of his efforts. Baird was instrumental in bringing to Leidy's Philadelphia laboratory the fruits of the Government survey collections. For many years Leidy, unable to accompany the western expeditions, was fully occupied with the fossils, which were never lacking in abundance. He was the American pioneer in paleontological research, describ- ing the extinct oreodonts, camels, rhinoceroses, and titanotheres that roamed the Miocene. His more than two hundred papers on paleontological subjects culmi- nated in a great work on the extinct mammalian fauna of Nebraska and Dakota (Leidy, 1869). This report includes a synopsis of the mammalian remains of North America. A fitting epitaph to this quiet and retiring scientist was given by Osborn, who praised him "as the last great naturalist in the world of the old type, who was able by both his capacity and training to cover the whole field of nature." Marsh and Cope completed the triumvirate of the early paleontologists, fol- lowing in the footsteps of Leidy. Independently wealthy. Marsh could muster his own expeditions. His graduate students at Yale accompanied the bone hunter on repeated expeditions to Colorado, Nebraska, Utah and Wyoming. Museums today display Marsh's prized collections of fossil horses, so valuable as a demonstration of evolution. These discoveries were among the finest of those made by Marsh. 670 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES VERNON BAILEY 1864-1942 ERNEST THOMPSON SETON 1860-1947 RUDOLPH MARTIN ANDERSON 1876- GERRIT S. MILLER, JR. 1869- HAMILTON: MAMMALOGY IN NORTH AMERICA 671 His ability as a field collector is reflected in the vast assemblage of mammalian fossils housed in Yale's Peabody Museum. A student of Leidy, Edward Drinker Cope continued the study of fossils. As a youngster, his first love was the reptiles, and the young naturalist made lasting contributions to our knowledge of salamanders and lizards. His major contribu- tions were made west of the Mississippi, while Cope was employed as vertebrate paleontologist of the U. S. Geological and Geographical Survey of the Territories. The contributions Cope made on the creodonts, canids, and felids were outstand- ing. His thousand-page volume on the vertebrates of the Tertiary Formations of the West includes accounts of 350 species, 90 percent of which the author had described (Cope, 1884). The collecting of fossil remains is a slow and tedious process. Yet even greater effort must be employed in the museum when reconstructing the fruits of expedi- tions. Following the period of western exploration, the study of collections con- sisted in establishing the lineage of the families, orders, and classes. Many groups have been collected which provide a panoramic view of lineal descent. Among paleontologists of the present century, Henry Fairfield Osborn must receive spe- cial recognition. He was a rare combination of scientist, teacher, and adminis- trator. From 1877, when he commenced paleontological research at Princeton, until his death in 1935, Professor Osborn published nearly a thousand articles and memoirs. Among his best known works are The Age of Mammals (1910) , Men of the Old Stone Age (1916) and The Titanotheres of Ancient Wyoming, Dakota and Nebraska (1929) . He somehow found time to write many popular articles and books, detailing the lives of creatures that lived in the past. A Princeton classmate of Osborn, "William B. Scott, contributed materially to the study of early mammals. His History of Land 3Iammals in the Western Hemisphere (1913), while designed primarily for lay readers, is of considerable service to the professional mammalogist. It is difficult to single out individuals who have made lasting contributions in any field of science without creating injustices. The names of Edwin H. Colbert, William K. Gregory, Claude W. Hibbard, Remington Kellogg, William D. Matthew, John C. Merriam, George G. Simpson, Ruben A. Stirton, Chester Stock, Horace E. AVood, 2nd, and Jacob L. Wortman merit especial notice for their substantial reports on fossil mammals. Of these, Simpson has made particu- larly noteworthy contributions in recent years. The Growth of Literature on Mammals With the development of mammalogy in North America, it was apparent that many works would appear dealing with this group. Mention has been made of the Quadrupeds of North America. This stellar contribution was a model for its time. Even today, serious students of mammal habits consult the three volumes, for there is a wealth of information that is remarkable for the years in which they appeared. It is probable that if the authors had used a model life history outline, as we know of such today, their immense background of knowledge would have resulted in an even more lasting contribution. Audubon and Bachman did less credit to themselves and the animals they discussed than they might otherwise 672 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES have done. The Quadrupeds is, nevertheless, an enduring monument to these two men. Robert Kennicott, dead in the Arctic at thirty years, was a disciple of Baird. When only twenty-two years of age, he published a report on the mammals of Illinois, which includes substantial information on many common species (Ken- nicott, 1858). The monumental Pacific Railroad report of Baird, who knew only 220 living kinds of North American mammals, was, in some respects, a standard until the time of Merriam. Between the two decades that separated the productive efforts of these two men, an amazing book appeared. In 1876, when only twenty-five years old, David Starr Jordan, then a young teacher in Wisconsin, published his A Manual of the Vertebrates of Eastern United States. It was, and still is, widely used, although now out of print. Jordan's Manual was the bible of many an early naturalist, for it included simple keys and brief descriptions of all of the known vertebrates occurring from the Atlantic coast to Iowa and south to North Caro- line. Primarily an ichthyologist, Jordan had received a good basic training in the vertebrates at Cornell University. The unparalleled publications of Merriam had set the style in the latter part of the nineteenth century. His Mammals of the Adirondaks, published in book form in 1884, remains a classic and a model for studies yet to come. We must not disregard the influence of Trouessart, whose Catalogue 3Iammalium (1897-1905) is a concise review of the then known mammals of the world. During the next decade the important American publications on mammals were largely confined to the reports of the Biological Survey. The North Ameri- can Fauna series appeared regularly, containing much on the systematics of our native species. To detail the many excellent reports on mammals that have made their ap- pearance in the past twenty-five years would be tiresome to the reader and serve no useful purpose. With what one writer might consider the highlights of achieve- ment in the field of literature, a score stand ready to disagree. Subject matter and the major references will be listed, if only to give an index to the breadth of the subject. The inquisitive reader will find, in the documentations of the studies referred to below, the more important references that attempt to cover the subject. General reference works on mammals are notable for their paucity. Flower and Lydekker's Mammals Living and Extiyict, published in 1891, and Beddard's classic Mammalia in the Cambridge Natural History Series are inclusive accounts of the mammals of the world. Surely these volumes, together with Weber's Die Saugetiere, appearing in 1904, may be considered outstanding. The excellent but smaller volumes of Angel Cabrera Manual de Mastozoologia and Dr. F. Bourliere's Vie et Moeurs des Mammiferes stress the ecological approach. E. W. Nelson's popular account of North American mammals had a salutary effect on the study of our native species. Published by the National Geographic Society in 1916 and 1918, this report was embellished by the peerleess artistry of Louis Agassiz Fuertes. A decade before the appearance of these studies, the more serious student of mammalogy was treated to W. D. Scott's History of Land Mammals in the Western Hemisphere. In 1929, Ernest Thompson Seton's Lives of Game Animals appeared. In his final words of the preface, he says : HAMILTON: MAMMALOGY IN NORTH AMERICA 673 ... I do not consider that I am offering even a fragmentary presentation of the final truth that is coming. This I feel — that I am merely assembling tools, and some day a great man will come, and with these tools construct a telescope that shall surely reveal to us the vision that the world is awaiting. Those of us who acknowledge Setoii's gift for writing, his industry, and his impressive stature as a field naturalist will not long forget his zeal and ambitions. His ability to portray the animals as he saw them has seldom been surpassed. Seton's final effort, indeed his life work, was directed to the Lives of Game Ani- mals, abetted by President T. R. Eoosevelt. In this fine study, the value of which will long be felt, Seton made his greatest contribution in a singular manner. He liberally quotes the sources unavailable to many of the present generation. Well documented, the volumes indicate the sources he searched so assiduously, such as Forest and Stream, the published journals of the older naturalists, and other re- ports that are often hard to come by. Diligent search in any sizable library will find the old notes but Seton brought them together. While one may read a dozen pages without learning much that is new, the fascinating manner in which Seton put them down will long be remembered. Many who read Seton's account of a species consider that what he did not record must be new. On the contrary, one can read pages without end in the Lives and find that the study of any one species is yet undeveloped. Seton's volumes on the game animals are a beginning. He amassed the data that have helped us all, but the work is an unfinished report, as Seton knew. More recently, the Mammals of North America by Victor Cahalane has pro- vided a wealth of information. His lucid accounts are detailed and provide a ready source of information on our native species. The volumes by Francis Harper Extinct and Yanisking Mammals of the Old World and the late Glover M. Allen's Extinct and Vanishing Mammals of the Western Hemisphere, published in 1945 and 1942 respectively by the American Committee for International Wildlife Protection, are models of inclusive but concise reports, so thoroughly documented that any biologist can ill afford to pass them by without study. The American Midland Naturalist, Ecology, and Ecological Monographs, Journal of Wildlife Mcmagement, the several AVistar Institute journals, and the publications of the state colleges and universities are rich in mammal lore. The many state academies have reports that are of interest to the mammalogist. The result of the monumental effort of Gerrit S. Miller, Jr., in compiling the List of North American Recent Mammals, appeared in 1924. This indeed was the crowning effort to a lifetime of research. Included are synonyms, type localities, and usually the range of all the known mammals inhabiting the area from Panama to Greenland. A revision of this important work is in press, Dr. Remington Kel- logg assisting Miller in the task. A more recent check list is that of Anderson (1946). In his account of Cana- dian recent mammals, Dr. R. M. Anderson, dean of Canadian mammalogists, compiled a lifetime study of the mammals in the provinces north of our border. His knowledge of Canadian mammals is evident in this report. The biology of any animal revolves, of necessity, around two major points. It must eat to live and, second, it must reproduce to perpetuate its kind. The com- prehensive story of reproduction is brought fully to date by Asdell's Patterns of Mammalian Reproduction, published in 1946, in which the author collated most 674 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES of the data bearing on the reproductive behavior of wild as well as domestic mammals. An excellent summary of the economic relations of mammals has been com- piled by Henderson and Craig (1932). This book includes a wealth of data on the practical aspects of mammals, particularly as related to man. The references on the dietary of wild mammals are quite complete. The volume is thoroughly documented. Probably more people have been attracted to the many fields of natural history by pocket guides than from any other source. Mammal books have not kept pace with allied fields in this respect. Bird guides, and good ones too, are without end; we have pocket editions of books that are aids in determining plants, insects, shells, and the like. A handy guide for the identification of mammals is another matter. Many species have subtle differences which are hard to see, let alone differentiate. Anthony's Field Book of North American Mmmnals, published in 1928, established the amateur's interest in mammals. Covering all of North America, Anthony included descriptions of species and their races, some maps indicating present known ranges, and some figures that were considered helpful for identification. It is a valued contribution to mammalogy. As this report is being written, yet another small book appears. A Field Guide to the Mammals, by W. H. Burt and Richard Grossenheider, is a specific example of the trend in American natural history. A splendid book, embellished with no end of colored plates, maps, and figures of tracks, it will serve as a model for years to come. Pocket books on natural history have undoubtedly brought many amateurs into specific fields of study, and many of these naturalists have made substantial contributions to our knowledge. Animals are no respecters of political boundaries. Yet the dictates of man all too often indicate that faunal surveys shall be made within a single state or province. Hence political lines, rather than natural boundaries, often limit the reports of these faunal studies. Many state reports on mammals have appeared. Among these, special mention must be made of Lyons' Mammals of Indiana, Ver- non Bailey's Mammals of New Mexico and The Mammals and Life Zones of Ore- gon, W. B. Davis's The Recent Mammals of Idaho, Burt's Mammals of Michigan, E. R. Hall's Mammals of Nevada, Dalquest's Mamynals of Washington and the comprehensive two-volume Fur-Bearing Mammals of California, by Grinnell, Dixon, and Linsdale. The account of a smaller region of a state, embracing a natural unit, is that of Harper (1927). This model report is one of the best local studies that has yet appeared. An excellent summary of the development of the classification of mammals from Aristotle to Weber has been recorded by Gregory (1910). Simpson (1945) adequately summarizes the works that have influenced the development of mam- malian classification. Some universities fortunately have their own publications and can thus pro- vide an outlet for substantial reports. Among these several institutions, most notable are the University of California Publications in Zoology, the University of Michigan's JMiscellaneous Publications in Zoology and the University of Kansas publications. The Wildlife Review, a mimeographed bulletin designed for the abstraction of articles bearing on wildlife management, first appeared in September, 1935. HAMILTON: MAMMALOGY IN NORTH AMERICA 675 It has served a real need of the legion engaged in this field. In the 73 issues that have appeared to date much mammal research has been summarized. The review- is far more inclusive than the title indicates. The American Wildlife Institute has sponsored reports of monographic scope on wolves, coyotes, and the puma. We may soon look for a treatise on deer. The Growth of Mammal Collections Early collections of mammals in the state cabinets and lyceums of natural history were notable only for their paucity. A century ago, the larger of these were owned by private collectors. With the growth of the large museums, many of the private collections were donated, sold, or bequeathed to the museums. In earlier days, most were displayed as mounted specimens, and emphasis was given the larger or more striking species. Since the primary function of a great museum is to promote research, it is apparent that large collections of the many species in a convenient form for study must be available to the specialist. For every mounted specimen in the showcases of the larger museums, usually more than a score and often hundreds are housed in the mammal collections reserved for study. Figure 1. The cyclone trap, and its later refinement into the snapback trap as we know it today, made modern mammalogy possible. A few dollars provides the collector with sufficient traps to make a survey of any region possible. The smaller trap has taken a Zapus, while the Museum Special holds a Condylura. Few inventions have been so instru- mental in furthering the growth and promotion of a specialized field in natural history. 676 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES We have seen the revolutionary change brought about by the invention of the snap-back trap. This little device, designed to take the smaller species, was responsible, within a few years after its appearance, for the hundredfold increase in the size of mammal collections. The mammal collection of the Fish and Wildlife Service (more familiarly known as the Biological Survey) is limited to recent North and Middle America species, of which it has the largest representation of any collection in the world. Included in this great collection is the type of the smallest North American mam- mal, Microsorex hoyi winnemana, a tiny shrew weighing less than a dime. This elfm creature was collected by Edward A. Preble on the Potomac shoreline, almost within sight of the building where it is now housed. The collection also includes the type of the largest of all existing carnivores, TJrsus tniddendorfi, collected on Kodiak Island, Alaska. Of the valued types, the survey collections contain 1,313, nearly half of the species and subspecies of North American mammals that are known today. These collections, like others, are indispensable in connection with the administration of wildlife, and are the basis for distributional, taxonomic, and identification studies. On June 30, 1952, this collection contained 146,237 catalogued specimens. The United States National Museum collection contains mammals from all parts of the world. In recent years, about 2,000 specimens have been added an- nually. This collection now (1952) has 110,824 specimens. Major collections have been received in the past ten years from Alaska, the Canadian Arctic, Labrador, Costa Rica, Panama, and Colombia in the New World, and from Egypt, Sudan, Japan, Korea, Formosa, Philippines, Burma, Nepal, Siam, Borneo, Australia, and nearly all the island groups of the Pacific. The American IMuseum of Natural History mammal collections contain more than 130,000 specimens. The many expeditions to South America, Asia, Africa, Madagascar, Australia, and Oceania have resulted in the discovery of many new species. Well over 800 types are represented in the museum. More than half of the collection is composed of North American specimens. In 1940, the Museum of Vertebrate Zoology, University of California, had in its collections slightly more than 100,000 skins. These were primarily representatives of the Pacific Coast, from Alaska to Lower California, the Great Basin, northern Mexico, and Salvador. This is a remarkably large collection for a university museum, more than twice the number contained in the University of Michigan, which may be considered the second largest mammal collection owned by an educational institution. The increasing interest in mammals is reflected in the ever growing collections, both private and public. A survey of the existing North American collections was made by A. B. Howell in 1923. A comparison of the survey made by Doutt et al. (1945) and Howell's earlier study reveals many interesting changes in the twenty-year span. In this short period the number of specimens in collections more than doubled. The number of private collections had increased two and a half times, while public collections were almost five times as common as in 1923. Doutt's report lists 939,483 specimens in United States and Canadian collections, whereas only 410,239 specimens were recorded by Howell in 1923. Since 1943 even greater strides have been made. The National Museum collection is increas- ing by 2,000 specimens a year, while more than 8,000 specimens were added to the Biological Survey collections in the past nine years. The University of Michigan HAMILTON: MAMMALOGY IN NORTH AMERICA 677 collections have increased correspondingly, approximately 10,000 specimens being added since Doutt s survey in 1943. Smaller collections have increased accord- ingly. The mammal collection at Cornell University has doubled in the past nine years. It is quite likely that other museums have added materially to their collections in the past decade. The Conservation of Mammals Early historians have left us with a record of the abundance of native mam- mals a century ago. The once widespread distribution of the big game species and their incredible numbers, even into the latter part of the nineteenth century, has been faithfully catalogued. The primitive population of the bison has been placed as high as 60,000,000, a figure which is probably extravagant. The prong- horned antelope, native only to western North America and tj^pically a resident of the Great Plains, probably rivaled the bison multitudes. In the middle of the last century, the lordly American elk or wapiti roamed through eastern forests from Quebec to Georgia. The whaling industry flourished, bringing riches to the adventurous sea captains and their hardy crews. The pelts of fur animals were much in demand, prompting hardy trappers and traders to invade the uncharted wilderness in quest of a harvest. Many eastern towns, rivers, and lakes have taken their names from the beaver, substantial evidence of its widespread distribution during the past century. The eventual decline and near extirpation of many of our larger mammals cannot be laid to any single cause. Insatiable greed and reckless slaughter by man with no thought to the future was surely one of the major causes of this decline. To be sure, the western plains could not support the livestock industry, the rolling miles of wheat, and the hordes of buffalo. These great hoofed creatures are now reduced to a few thousand semidomesticated animals herded on Federal and private reservations. A free herd in the Wood Buffalo National Park of Canada may be considered the only truly wild bison existing in North America. Except for sporadic introductions, elk have disappeared from the East, and are now largely restricted to the mountain country of the West. . By the turn of the century, beaver had all but disappeared from the eastern forests. Market hunting had been a notable instrument in the reduction of the deer. Settlement of the country encouraged large-scale agricultural operations, while the trans- continental railroad provided a ready means of getting wild meat to the eastern markets. The continuing demand for hides and pelts resulted in further inroads on our native mammals. Small wonder that state and Federal authorities and all interested in our natural resources were alarmed at the appalling destruction. Their concern is no less marked for species that today appear headed the way of the bison. Less than a half-century ago the great Merriam elk, unable to com- pete with cattle on the overgrazed range and susceptible to hunting pressure, disappeared forever. Is the end at hand for the little Key deer, Odocoileus vir- ginianus clavium t Inhabiting an area only 17 miles long and 15 miles wide, this diminutive creature has the smallest range of any deer in the world. A full grown buck of this elfin race stands but 25 inches at the shoulder and weighs little more than 30 pounds. The entire population of the Florida Keys was esti- mated at 57 individuals in January, 1952. 678 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES The death knell has sounded for many North American mammals. The pic- ture, however dark, is not quite one of such despair as many like to indicate. Much of the destruction of this natural heritage has been due to ignorance and thought- lessness. It will be appropriate to consider some of our native mammals whose threatened extinction a few decades ago was of grave concern to the American public. Few stories are more impressive in conservation history than that of our fur seals. The ravishment of the great herds had been carried on for nearly a century and a half when the Russian navigator, Gerassim Pribiloff, discovered, in 1786, the islands that bear his name. In that year, probably 4,000,000 seals occupied the rocky shores during the spring and summer months. Pelagic hunting by fish- ermen of Canada, the United States, and Japan had resulted in such reduction and waste that by 1910 not more than 130,000 animals remained of the former millions. In 1911, a treaty between Russia, Japan, Great Britain, and the United States put an end to pelagic sealing, and our country, owning the islands, as- sumed management of all sealing operations. A quarter-century later, the herds totaled 3,600,000 animals. Fish and Wildlife Service personnel cooperate with the Fouke Fur Company in handling the seal harvest. The animals are driven from their rocky hauling grounds to the flat tundra, where groups of immature males are cut out and the remainder allowed to return to the sea. The number annually killed is based on the size of the herd. The increasing returns from the sale of pelts (60,000 to 70,000 annually) and by-products has provided the gov- ernment with a growing profit and at the same time assured a livelihood to the natives of these lonely shores. The exploitation of the great whales followed a pattern of many another natural resource. In the early years of the past century, whaling was confined largely to coastal waters. Later the whalers ventured on all of the oceans of the world; the United States owes much to the intrepidity and fearlessness of the hardy whaling masters who first carried the American flag into new and little explored corners of the world. The decline in the number of whales has been evi- dent for many years, but improved methods of hunting and handling the catch of whales and the utilization of by-products make whaling still profitable to those engaged in the industry. The fleet of vessels and floating refineries returning from the South Seas in 1930 brought the largest cargoes of sperm oil ever loaded. These whales were located and reported by wireless-equipped aircraft and killed by electric harpoons. It is fortunate that the leaders in the whaling trade are cooperating in an effort to obtain data on these cetaceans which will be helpful in evaluating the biological factors involved. At the turn of the century, whalers began operating in the Antarctic Ocean, the last great unexploited area. In the early 'thirties the League of Nations called together a committee to consider international regulation of the whaling industry. Since this action, several international conventions have set forth regulations for whaling, the first in 1932 and another in 1937, upon which, with subsequent protocols and agreements, the present whaling regulations are chiefly based. These regulations prescribe seasons for whaling, establish the minimum legal size of each species, and prohibit the killing of females accompanied by calves, and of any whales of certain species. The regulations also require the fullest possible HAMILTON: MAMMALOGY IN NORTH AMERICA 679 use of each whale taken. The participating nations, which include most of the important whaling countries, share responsibility for enforcement (Carson, 1948). The history of the beaver in North America follows a pattern well known to conservationists. At one time it was widespread and abundant in the east but trapping pressure for the valued pelts brought it virtually to the brink of exter- mination. By 1900 New York and the New England states could boast of only a few dozen. Introductions of a few here and there resulted in an astonishing increase. In the early 'twenties, increasing complaints of damage indicated all too well the success of these introductions. The beaver is now actually a pest in many of the regions where it was a rarity a half-century ago. Through the lElood- ing of valuable timberland this big rodent may actually prove a nuisance. The white-tailed deer is another striking example of a species that became so scarce in the early part of the present century that Easterners considered it no longer of significance as a game species. Introductions and closed seasons have now made this fine animal abundant in the East. Its unprecedented increase in recent years has been cause for much concern among agriculturists, for deer depredation in orchards and to crops is of no mean consequence. Some Practical Considerations With increasing human populations, it was apparent that the wildlife of North America would play an ever more important role. Environmental changes wrought by man resulted in far-reaching effects. Lumbering operations destroyed habitat for the moose and bear, while it created a more desirable habitat for the cottontail and fox. The resultant farmlands and second-growth timber provide a more suitable environment for many species that shun the solid stands of timber. Destruction of grasslands on the western prairies increased competition between rodents and livestock for the range. These changes have been reflected in many ways. It is difficult, often impossible, to assess an animal in the economic ledger. The common field mouse plays a useful part in the economy of nature when it occupys waste lands. Here it provides food for a host of predators, transforming grass into fur coats. It may act as a buffer against predation on more desirable species. In the orchards and grain fields, its ravages are measurable; here it must be classed as a pest of the first order. We acknowledge the usefulness of the beaver in impounding waters and preventing rapid run-off. Its value in the past and present as a fur-bearer will not be denied. When the big rodent kills extensive tracts of valuable timber through flooding, or disrupts a water supply through interference with the normal water level, then we must take steps to control the animal. The cottontail rabbit is hailed as our primary game animal in the eastern states, yet its depredations in the orchard or garden are often severe. It must now be apparent that a decision regarding the economic value of a species is difficult, indeed, often impossible, in the light of our present knowledge. Judg- ment of any species must take into consideration many factors, two of the most important being time and place. We may consider several categories, when attempting a critical judgment of the economic worth of a species. 680 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES The esthetic value of many mammals cannot be denied. Many thousands of tourists visit our National Parks annually. "Wild animals, be they elk, bear, wild sheep, or the teeming populations of smaller species, share with the geysers, waterfalls, and great forests the interest of the people. Summer visitors to our eastern parks delight in the sight of a beaver dam, and drive many miles in the hope of catching a glimpse of a doe with her fawns on some wilderness meadow. Residents of Vilas County, Wisconsin, realizing the recreational value of wildlife to the tourist, posted thousands of acres against hunting. These people realized that deer were a greater asset to them alive and brought a larger reward through summer trade than nonresident hunters could possibly do. In 1946, over 21,000,- 000 people visited the 160 acres of the National Park system. They came to see our native mammals as well as the other natural wonders. Perhaps more tangible values are to be found in the hunting and trapping of game and fur-bearing species. An increasing number of hunters take to the fields each year. It is fortunate for them that many game species have shown a like increase in numbers. The exploitation of our fur resources is woven inextricably with the settlement of the great Mississippi Basin and the West. We have seen that the resultant changes in environment have been responsible for the decline of many species, while others have increased wherever man has partly cleared the forests and farmed the land. Some species are adaptable and can thrive in arable lands, whereas others depend upon wilderness areas. Trapping is big business and provides a partial livelihood to many thousands of Americans. In the early 'forties, trapper income was estimated at no less than $100,000,000 annually. While this may appear to be a relatively small figure in so far as products of the land are concerned, the return is very substantial. The money is distributed among the low income group and at a season when a cash crop is most needed. The fur industry is a huge one, employing many people who are directly depend- ent on this great resource. Except for the'muskrat and the beaver, we know less about the habits and needs of our fur animals, than we do of the game species. This lack of knowledge may be attributed to several factors. Most important, perhaps, has been the almost universal belief that fur-bearers and vermin are synonymous. This has been particularly true of the weasel, mink, skunk, fox, and other carnivores. The apathy of state game officials has been marked. Fur animals have brought little or no revenue to the state treasuries, hence research on, and legislation for, this valuable resource has not until recently received the attention it merits. The annual loss to crops, forage, and forests occasioned by our native mammals is a very real one. Bell (1921) has. placed this monetary loss at $300,000,000. By far the larger share of this loss may be levied against pocket gophers, ground squirrels, field mice, cottontails andjackrabbits, with cotton rats, porcupines, woodchucks, moles, and other species adding to this destruction. It is a well known fact that wild mammals may transmit virulent diseases to man and his livestock. The study of the diseases of wild mammals is still in its infancy; man and his domestic animals frequently contract these diseases. AVhen outbreaks of rabies, tick fever, or endemic typhus break out among feral species, it has been found necessary, often at considerable expense, to conduct extensive campaigns against these animals. Such wholesale slaughter is regrettable but inevitable when con- HAMILTON: MAMMALOGY IN NORTH AMERICA 681 siderable monetary loss or a threatened human pandemic appears miminent. When the 1924 outbreak of hoof-and-mouth disease occurred in California, more than 22,000 deer were poisoned on the Stanislaus National Forest of California. The disease was checked, and the deer soon regained their former abundance. Mammalian Eesearch It was inevitable that research studies should emphasize the species which are of economic significance. Following the early trend of systematic mammalogy, when species were described, their distribution plotted, and their practical sig- nificance determined, emphasis was directed toward the acquisition of detailed knowledge concerning individual species. With the essential features of the dis- tribution of most of our commoner species mapped, the main categories in the life histories have been catalogued, if only in a brief, superficial manner. To be sure, this advance in our knowledge of the rich North American mammalian fauna has not kept pace with the more numerous bird species, but the reason is quite apparent. In the early years of field investigation, emphasis was placed on regional lists, annotated with brief accounts of the habits of the included species. These reports followed the pattern of some of the early North American Fauna series. In these Figure 2. By their numbers alone, the teeming hordes of ground squirrels in west- ern North America provide an unparalleled opportunity for research. Behavior, activity and population studies, to cite a few, are indicated by the abundance of these little mammals. The golden-mantled ground squirrel, Citellus lateralis, was photographed in Estes Park, Colorado, on July 1, 1941. 682 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES reports, emphasis was placed on the distribution of species, but much new material on habits was included. The detailed studies currently being conducted on the larger mammals are noteworthy. Federal and state funds have been made available for these researches. In 1949, thirty-four states were engaged in deer research alone. While duplication of effort is inevitable in such widespread investigations, the end results may well justify these many studies. Many notable contributions have been made on the life history of a single species, but in the present brief summary, mention can be made of only a few. Paul Errington's muskrat studies are of major import. His principal objective has been to fathom the rules of order governing the distribution and maintenance of populations in different types of habitat. The studies of Lee R. Dice, of the University of Michigan, and his students, notably W. Frank Blair, on speciation in Peromyscus indicate the value of long-time research on a single genus. The laboratory studies of Dice support the keen taxonomic judgment of the late Wil- fred Osgood, whose revision of the genus is an unparalleled systematic study. The present trend is not so much an effort to catalogue the details of a par- ticular species, but rather to attempt interpretations in the light of relationships. We must now inquire into the "why" rather than solely occupying ourselves with observed facts. A classic account of field observation has been the documentation of the behavior of the red deer (Darling, 1937). This report should be studied by all naturalists. The publication of Charles Elton's Animal Ecology in 1927 lent great impetus to the study of population dynamics. Presently a score of American investigators are following his leadership in this important field. The development of banding or marking animals so that they might be recognized when recaptured has been of inestimable value in determining many biological features. This sub- ject, together with territorialism, was first studied among the birds by H. E. Howard. The initial studies on mammals in this field were made by W. H. Burt, of the University of Michigan. Systematic mammalogy has undergone a marked change in the century under review. Earlier taxonomists were content to describe a new species in a few hundred words, expressing little concern for the relationships that existed between the new form and its close relatives. The concept of modern taxonomy rests in an expression of relationship, based on the study of large series and firsthand knowl- edge of the habitat in which the species lives. An excellent example is the recent study of the harvest mice by Hooper (1952) . The American Society of Mammalogists The early interest in ornithology and entomology may, in a measure, be at- tributed to several factors. Their subjects attract the eye, they are everywhere to be seen, and their great variety and the relative ease of collecting them attracts the young naturalist. The making of collections and the exchange of specimens has had a stimulating effect on the development of natural history in North America. We have seen how the development of the snap-back trap had a salu- tary effect on the growth of mammalogy. The time was ripe for the organization of a society, the function of which would be to encourage interest in mammals and provide a means for the publication of original research. Botanical, entomo- logical, and ornithological societies were flourishing in the early years of the HAMILTON: MAMMALOGY IN NORTH AMERICA 683 present century, but there was yet no organization devoted solely to the study of mammals. The American Society of Mammalogists was founded at Washington, D. C, on April 3, 1919. When a call was issued for a meeting at the U. S. National Museum on that date, sixty persons from many parts of the United States and Canada were present for the initial session. Plans for the society were perfected, officers elected, committees formed, and by-laws and rules were adopted. The objects of the infant society were declared to be "the promotion of the interests of mam- malogy by holding meetings, issuing a serial or other publications, aiding research, and engaging in such other activities as may be deemed expedient." Systematic work, life history and habits of mammals, evolution, paleontology, anatomy, and every phase of popular and technical mammalogy were to be within the scope of the society and its publication. One of the declared objects of the society was to be the publication of the Journal of Mammalogy. This publication was planned to be indispensable to all workers in every branch of mammalogy and of value to every person interested in mammals, be he systematist, paleontologist, anat- omist, museum or zoological garden man, sportsman, big game hunter, or just plain naturalist. How well the aim has been fulfilled is attested by the breadth and wide scope of articles that have appeared in the thirty-three volumes of this quarterly. The first officers and directors are a virtual roster of the great names in American mammalogy thirty years ago. Merriam was honored with the presidency, a fitting tribute to his lifetime contributions in the field. E. W. Nelson and Wilfred H. Osgood were vice-presidents. Hartley H. T. Jackson the corresponding secretary, Walter P. Taylor, treasurer, and Ned Hollister the editor. Members of the coun- cil (now known as directors) included Glover M. Allen, Eudolph M. Anderson, Joseph Grinnell, Marcus W. Lyon, Jr., W. D. Matthew, John C. Merriam, T. S. Palmer, Edward A. Preble, and Witmer Stone. Of this group, Nelson, Osgood, Matthew, Allen, Stone, Lyon, Grinnell, Jackson, and Taylor all were elected to the presidency of the society in recognition of their contributions to mammalogy. The society has published several monographs, including the Anatomy of the Wood Rat by A. B. Howell, The Beaver: Its Work and Ways by E. R. Warren, and Animal Life of the Carlshad Caverns by Vernon Bailey. The Society has a current membership of more than 1,350 individuals. The Pacific Northwest Bird and Mammal Society was founded January 7, 1920. One of the objects of the society was to promote interest in the scientific study of birds and mammals within the region mentioned. The Murrelet, official publication of the society, is published triannually. Except the American Society of Mammalogists, this is the only organization in the United States which ex- pressly designates the professional field of mammalogy as one of its primary aims. In Germany, the Deutschen Gelleschaft fiir Saugetierkunde, organized in 1926, publishes the Zeitschrift fiir Saugetierkunde. The French journal Mammalia has gone through sixteen volumes. In content, these two important journals follow the leadership of the Journal of Mammalogy. We would be remiss, indeed, if no mention were made of the many other societies that have not only professed an interest in mammals, but, by militant effort, have helped further interest in our native species. The National Audubon Society, aware of the plight of many of our rarer mammals, has been increasingly 684 ^ CENTURY Of PROGRESS IN THE NATURAL SCIENCES concerned with conservation problems. Among Federal agencies, the Soil Con- servation Service, the Bureau of Animal Industry, the Bureau of Entomology and Plant Quarantine, the National Park Service, and the Forest Service all are interested in the several biological fields which include consideration of mammal life. In close cooperation with the Fish and Wildlife Service, they are directly responsible for research that often includes the study of mammals. Present Needs Surely it is evident that many pressing problems of utmost economic signifi- cance and academic interest have yet to be solved. Research on our native mam- mals, except within recent years, has not been extensive. It is lamentably true that problems dealing with mammals currently arise, some of great importance, that cannot be answered with authority. Training in mammalogy is as necessary to those who would consider it a profession as it is in other allied fields. A knowl- edge of botany, entomology, geology, mathematics, and kindred subjects must be considered a part of the training of the professional investigator. In earlier years, a few recognized masters guided the destiny of many an untrained youth into the field in which he later excelled. The professional mam- malogists of the past century were primarily trained in medicine, as were Mer- riam, Mearns, and Coues. Others had no formal training in the sciences. Vernon Bailey is an illustrious example of a self-taught naturalist who gave inspiration to the generation that followed. In the early part of the present century, few educational institutions were concerned with specialized courses in natural history. The classic instruction included anatomy, physiology, and embryology, for the prescribed curriculum was designed for premedical training. The influence of Agassiz, Jordan, and their disciples was destined to foster the study of living animals. In the training of a naturalist, be he interested in systematics, morphology or life histories, a sound biological basis is the best preparation. Few will deny this assumption. University courses in the natutral history of vertebrates, in which the study of mammals was included, were given a half-century ago. Notable among the institutions that gave special instruction in mammalogy in the early years of the present century should be mentioned the University of California, Cornell University, and the University of Michigan. Presently thirty or more colleges give courses that deal with mammals, while many others offer instruction in vertebrate natural history. Mammal studies are emphasized in many schools that include wildlife courses in their curriculum. These, of necessity, differ widely in the various institutions where such instruction is offered. Ever since the first World AVar, when attention was focused on the alarming state of our natural resources, including wildlife, the growth of both Federal and state agencies concerned with the management and conservation of these resources has been marked. Instruction in the universities has kept apace with the increased demand for trained personnel to fill the positions in this expand- ing field. While mammalogy is only a small part of the wildlife field, there is a constant demand for individuals with specialized training in this subject. In government, state, and educational institutions, the training requirements are rigorous and selective. Only the ablest candidates are assured of a position. HAMILTON: MAMMALOGY IN NORTH AMERICA 685 Since these are career positions, promotions are usually slow, and salaries are not comparable to those of the other professions. Positions in the field of mammalogy are not numerous, and the prospective student planning a career in this branch of zoology should examine carefully the opportunities before embarking on a specialized course. Basic training in the natural sciences, including mathematics, geology, chemistry, physics, and the usual undergraduate courses in biology, are a primary requisite to advanced study. Graduate study is desirable, but not of paramount importance if the individual has a broad concept of the field. This is usually acquired during the early years and follows the usual pattern of collecting and an interest in a particular group, be it plant or animal. In an interesting report on this subject Miller (1928) stated: As now used, the term mammalogy applies primarily to what is known as the sys- tematic study of mammals, the main object of which is to find out exactly how many kinds of mammals there are in the world, exactly where each kind lives, and exactly what are the relationships of these creatures to each other and to their predecessors now gone from the ranks of living things. Included are systematics, distribution, and paleontology. Miller's definition of the science of mammalogy does not consider the economics, ecology, and life history of the mammals. The present trend in research is partially evident in a review of the past eleven issues of the Journal of Mammalogy. Of 118 major articles, more than half deal with the life history or habits of mammals, morph- ology accounts for 10 per cent, and systematics and distribution 8.5 per cent respectively. Since there appear to be somewhat fewer publication sources for the accounts of habits than for those on systematics, this evaluation does not give an accurate trend in mammal studies currently in progress. It does, how- ever, suggest the broad interests of the investigators presently engaged in mammal research. The advance in our knowledge of systematics and distribution has been particularly gratifying. Nevertheless, a promising field of investigation awaits those who are willing to spend long hours afield, collecting and observing in their natural haunts almost any species of North American mammal. The increas- ing number of young men and women that are being attracted to the study of mammals will surely have a salutary effect on the progress of mammalogy in North America. LITERATURE CITED Allen, Glover M. 1942. Extinct and Vanishing Mammals of the Western Hemisphere with the Marine Species of all the Oceans. Spec. Publ. No. 11. vii-xv + 620 pp.; illus. New York: American Comm. for International Wildlife Protection. Anderson, Rudolph Martin 1946. Catalogue of Canadian recent mammals. Nat. Mus. Canada. Bull. 102, pp. 1-238. Anthony, Harold E. 1928. Field Book of North American Mammals, iii-xxvi + 674 pp. 150 figs. 48 pis. New York: G. P. Putnams Sons. 686 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES ASDELL, S. A. 1946. Patterns of Mammalian Reproduction, v-x + 437 pp. illus. Ithaca: Comstock Publ. Co. Audubon, John James and John Bachman 1845-1854. The Viviparous Quadrupeds of North America. Vol. 1: vii-xiv, 1-389; Vol. 2, 1-334; Vol. 3, i-iii, 1-348. New York: V. G. Audubon. Bailey, Vernon 1907. Wolves in Relation to Stock, Game and the National Forest Preserves. Forest Service Bull. No. 72. 1931. Mammals of New Mexico. 412 pp. illus. North American Fauna 53. Bur. Biol. Surv., U. S. Dept. Agric. Washington, D. C. 1936. The Mammals and Life Zones of Oregon. 416 pp., illus. North American Fauna 55. Bur. Biol. Surv., U. S. Dept. Agric. Washington, D. C. Baibd, Spencer Fullerton 1857. Mammals: General Report upon the Zoology of the Several Pacific Railroad Routes. Repts., Explorations and Surveys for Railroad Route from Mississippi River to Pacific Ocean. 8(pt. 1) :xix-xlvii, 1-757, pis. 17-60. Washington, D. C. Beddabd, Frank Evers 1902. Mammalia, v-xii + 605 pp., 285 figs. London: Macmillan Co. Bell, W. B. 1921. Death to rodents. Yearbook U. S. Dept. Agr. (1920), pp. 421-438, illus. BOURLIEKE, F. 1951. Vie et Moeurs des Mammiferes. 249 pp. illus. Paris: Payot. Burt, William Henry 1946. The Mammals of Michigan, vi-xv + 288 pp. illus. Ann Arbor: Univ. Michigan Press. Burt, William Henry and Richard P. Grossenheider 1952. A Field Guide to the Mammals, v-xxi + 200 pp. illus. Boston: Houghton Mifflin Co. Cabrera, angel 1922. Manual de Mastozoologia. vii-xi + 440 pp. illus. Madrid and Barcelona: Com- pania Anonima de Libreria. Cahalane, Victor H. 1947. Mammals of North America, vii-x + 682 pp. illus. New York: Macmillan Co. Carson, Rachel L. 1948. Guarding Our Wildlife Resources. Conservation in Action, No. 5, Fish and Wildlife Service. Washington, D. C. Cope, Edward Drinker 1884. The Vertebrata of the Tertiary Formations of the West. Book 1. Report of the U. S. Geological Survey of the Territories. (F. V. Hayden.) Vol. 3. xxxv + 1009 pp., 75 pis. Dalquest, Walter W. 1948, Mammals of Washington. Vol. 2:1-444, 140 figs. Univ. Kansas Publ. Mus. Nat. Hist. Darling, F. F. 1937. A Herd of Red Deer. 240 pp. Cambridge: Oxford University Press. HAMILTON: MAMMALOGY IN NORTH AMERICA ^37 Davis, William B. 1939. The Recent Mammals of Idaho. 400 pp. illus. Caldwell, Idaho: Caxton Printers. Dekay, James E. 1842. Zoology of New York. Pt. 1. Mammalia, vi-xii + 146 pp., 36 pis. Desmarest, a. G. 1820. Mammalogie ou description des especes. Paris. DouTT, J. Kenneth, A. Brazier Howell, and William B. Davis 1945. The Mammal Collections of North America. Journ. Mammal. 26:231-272. Flower, William Henry, and Richard Lydekker 1891. An Introduction to the Study of Mammals Living and Extinct, v-xvi + 763 pp., 375 figs. London: Adam and Charles Black. Godman, John D. 1826. American Natural History, or Mastology. 3 vols. Philadelphia: H. C. Carey and I. Lee. Gregory, William K. 1910. The Orders of Mammals. Bull. Amer. Mus. Nat. Hist. Vol. 85, 350 pp. Gbinnell, Hilda W. 1943. Bibliography of Clinton Hart Merriam. Journ. Mammal., 24:436-457. Grinnell, Joseph, Joseph S. Dixon, and Jean M. Linsdale 1937. Fur Bearing Mammals of California. Vol. 1, vii-xii + 375 pp. Vol. 2, vii-xiv + 400 pp. illus. Berkeley: University of California Press. Hall, E. Raymond 1946. Mammals of Nevada, v-xi + 710 pp. illus. Berkeley: Univ. Calif. Press. Harlan, Richard 1825. Fauna Americana, x, 11-318. Philadelphia: A. Finley. Harper, Francis 1927. Mammals of the Okefenokee Swamp region of Georgia. Proc. Boston Soc. Nat. Hist. Vol. 38, no. 7, pp. 191-396. Boston. 1945. Extinct and Vanishing Mammals of the Old World. Spec. Publ. No. 12. x-xv + 850 pp., 67 figs. New York: American Comm. for International Wildlife Protection. HENDER.SON, JUNIUS, and Eliserta L. Craig 1932. Economic Mammalogy, v-x + 397 pp. Baltimore: Chas. C. Thomas. Hooper, Emmet T. 1952. A systematic review of the harvest mice (genus ReithrodontoTnys) of Latin America. Misc. Publ. Mus. Zool. Univ. Mich. No. 77, pp. 1-225, illus. Jordan, David Starr 1929. Manual of the Vertebrates of the Eastern United States. Thirteenth Edit., vii-xxxi, 446 pp. illus. New York: World Book Co. Kellogg, Remington 1946. A century of progress in Smithsonian biology. Science, 104 (no. 2693) : 132-141. Kennicott, Robert 1858. The quadrupeds of Illinois injurious and beneficial to the farmer. Rept. Com- miss. Patents (1857), pp. 72-107. 688 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Leidy, Joseph 1869. The Extinct Mammalian Fauna of Dakota and Nebraska including an Account of Some Allied Forms from Other Localities, together with a Synopsis of the Mammalian Remains of North America. Journal of the Academy of Natural Sciences Philadelphia, ser. 2, Vol. 7, 472 pp., 30 pis. Philadelphia. Lyon, Marcus Waed, Jr. 1936. Mammals of Indiana. Amer. Midi. Nat. 17:1-384. Merriam, Clinton Hart 1884. The Mammals of the Adirondack Region. 316 pp. New York: Privately pub- lished. Merriam, C. Hart, and Leonhard Stejneger 1890. Results of a Biological Survey of the San Francisco Mountain Region and the Desert of the Little Colorado, Arizona. North American Fauna. No. 3. Bur. Biol. Surv., U. S. Dept. Agr. 136 pp., 14 pis., 5 (col.) maps. Miller, Gerrit S. Jr. 1924. List of North American Recent Mammals, 1923. U. S. Nat. Mus. Bull. 128. xvi + 673 pp. Washington, D. C. 1928. Mammalogy and the Smithsonian Institution. Ann. Rept. Smithson. Inst. Publ. 2981, pp. 391-411. Nelson, E. W. 1916. The Larger North American Mammals. Nat. Geog. Mag. vol. 30:385-472. 1918. Smaller Mammals of North America. Nat. Geog. Mag., vol. 33:371-493. OsBORN, Henry Fairfield 1910. The Age of Mammals in Europe, Asia and North America, xx + 635 pp. New York: Macmillan Co. 1916. Men of the Old Stone Age, Their Environment, Life and Art. xxvi + 545 pp. New York: C. Scribner & Sons. 1929. The Titanotheres of Ancient Wyoming, Dakota and Nebrasks. Monogr. U. S. Geol. Surv., No. 55, Vol. 1, pp. i-xxiv, 1-701, 42 pis. Vol. 2, pp. i-xi, 702-953, 193 pis. Osgood, Wilfred H. 1943. Clinton Hart Merriam— 1855-1942. Journ. Mammal., 24:421-436, 1 pi. Scott, William B. 1913. A History of Land Mammals in the Western Hemisphere, ix-xiv, 693 pp., 304 figs. New York: Macmillan Co. Simpson, George Gaylord 1945. The Principles of Classification and a Classification of Mammals. Bull. Amer. Mus. Nat. Hist. Vol. 85. xvi + 350 pp. Trouessart, Edouakd Louis 1897-1905. Catalogue mammalium tarn viventum guam fossilium. Six parts, 1469 pp. Berlin: R. Friedlander und Sohn. Weber, Max 1904, Die Saugetiere. Einfuhrung in die Anatomie und Systematik der recenten und fossilen Mammalia, xii + 866 pp., illus. Jena: Gustav Fischer. INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY FROM 1850 TO 1950 By CHARLES E. WEAVER For a proper understanding of the development of invertebrate paleontology and historical geology from 1850 to 1950 a review of the important trends in research during the first half of the nineteenth century is essential. The early contributions which laid the foundations of these sciences originated for the most part in Europe, although there was marked advance in North America be- tween 1830 and 1850. Invertebrate Paleontology Prior to 1850 The publication of the tenth edition of Sy sterna Naturae by Linnaeus in 1758 laid the foundation of modern systematic zoology and invertebrate paleontology. Over 4,200 different kinds of animal life were listed, briefly described, and clas- sified according to a binomial system in which each form was given a generic and specific name. Linnaeus considered a species as composed of individuals descended from ancestors with common morphological characters and held that each separate species possessed certain immutable characters which remained constant and were not subject to modification. Interbreeding was possible only among individuals of the same species. This concept of a species strongly infiu- enced contemporary students of organic life during the later years of the eight- eenth century and the earlier decades of the nineteenth. Among the more important contributors to the development of paleontological science following Linnaeus was Georges Cuvier of France. Although his investi- gations were largely confined to fossil vertebrates, the principles developed were applicable to the invertebrates also. His earlier work involved a study of the anatomy of fossil bones of elephants from the Paris Basin and emphasis was placed on the differences in the skeletons of living forms in the collections of the Paris museums. He called attention to the evolution of these organisms. The first quarter of the nineteenth century was devoted to a comparative study of the osteology of the fossil remains of amphibians, reptiles, and mammals from the Tertiary deposits of Europe and a comparison of these bones with those of living representatives. The results of these important studies appeared in a four-volume work first published in Paris in 1811-1812. The significance of this contribution was the establishment of the law of correlation of parts. According to this law all the different components of the skeleton of an organism are mor- phologically related and a modification of one part would present corresponding differences in the other correlated parts. Many new genera and species from the Upper Eocene of the Paris Basin were described. The importance of this new [689] 690 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES procedure in the investigation of fossil remains is evident in the character of published work throughout Europe during the second quarter of the nine- teenth century. Cuvier emphasized the occurrence of the more primitive forms of animal life in the older geological formations and pointed out that extinct genera lived at an earlier time than those of the Recent. However, he did not believe that the living organisms had originated through the anatomical modifications of the older forms. He considered that during the past history of the earth there had been many sudden and violent disturbances of the crust which had resulted in the submergence of vast areas of land and the reappearance from beneath the ocean's surface of vast islands and continental masses. Destruction of the faunas and floras accompanied these catastrophic movements. Following each catastrophe a new and more advanced biologic fauna came into existence, the last of these appearing about six thousand years ago. Translations of Cuvier's work into several languages accentuated discussion of the problem. The Cuvierian concept of immutability of species was opposed by several European naturalists who held that new species might arise by the gradual modification of pre-existing species as the result of changing environments. This became known as the theory of mutability in contrast to that of immutability as advocated by Cuvier. Among the earliest strong supporters of the concept of mutability of species was Lamarck, who developed the idea that the morphological characters of a species might be subject to modification from generation to generation when subjected to environmental stimuli and that such acquired characters could be inherited. He recognized the morphological differences of extinct species from a succession of epochs and compared them with species now living in nearby areas and thus pointed out the possibility of the correlation of formations by the use of fossils. His investigations were largely concerned with marine inverte- brate fossils from the Tertiary deposits of the Paris Basin, which were described (Lamarck, 1815-1822) in his important monograph on Natural History of In- vertebrate Animals. This contribution, devoted largely to fossil mollusks, played an important part in the foundation of scientific conchology. Although the La- marckian concepts were accepted by an increasing number of European investi- gators yet the Cuvierian ideas were strongly entrenched in scientific thinking even beyond the middle of the nineteenth century and to a considerable extent influenced the writings of d'Orbigny. Many significant contributions were made to the growing science of paleon- tology during the early half of the nineteenth century. These consisted largely of descriptions of fossil species and monographs of faunas from different for- mations of Europe and North America, together with catalogues containing named species. Among the important contributors were James Sowerby and his son, James de Carle Sowerby, E. F. Schlotheim, H. G. Bronn, G. A. Gold- fuss, A. d'Orbigny in Europe, and T. A. Conrad in North America. The mollusks of Great Britain were described and illustrated in a six-volume work by the Sowerbys published from 1812 to 1846. This work was of great value for further investigations in conchology and for a comparison of the Ter- tiary faunas of Great Britain with those of France and other parts of Europe. Contemporaneously in Germany Ernst von Schlotheim in 1820 published his work Die Petrefadenkunde, in which many invertebrate fossils were figured WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 691 and described according to the binomial nomenclature. This work was important for those who were later concerned with prolilems of taxonomy. The rapid ap- pearance of published descriptions and illustrations of fossils were followed in 1834-1838 by a summary of the known information concerning paleontology and stratigraphy in H. CI. Bronn's Lethaea Geognostica and his Index Palaeon- tologicus in 1848-1849. Tliese works were of fundamental importance to many of the investigations carried on during the middle of the nineteenth century. The Petre facta Germaniae of Goldfuss and Munster carries descriptions and il- lustrations of fossil echinoids, mollusks, corals, and sponges collected largely in Germany. The publication of d'Orbigny's Paleontologie frangaise began in 1840 and continued till 1855. It was his intention to include a description with illustrations of all the fossils found in France but it was confined largely to Jurassic and Cretaceous echinoids, brachiopods, gastropods, and cephalopods. In North America James Hall had published several monographs on the Paleozoic fossils of New York State but most of his contributions appeared dur- ing the second half of the century. Intensive investigation of Tertiary mollusks was initiated by T. A. Conrad in 1832-1833 in his work on the fossil shells of the North American Tertiary. The first Tertiary fossil collections made on the Pacific Coast were submitted to him for identification and age determination and initiated a series of investigations which have been in progress for over one hundred years. It is of interest to note that Conrad followed the Cuvierian con- cept of species and believed that the fauna of each period of geologic time suf- fered annihilation as the result of climatic and other environmental changes, new faunas being developed in the following period. By the middle of the nineteenth century the Cuverian concept that the faunas of each geologic period had no species in common with those which preceded and followed it was gradually abandoned as information became available that tran- sitional genera and species partially filled the gaps and that the time span for each showed great variations. Although d'Orbigny, Agassiz, and others still supported in varying degrees the views of Cuvier, the evidence presented by Bronn that the faunas of each period resulted from the modification of species of the preceding period, together with the uniformitarian ideas advocated by Lyell concerning earth history, laid the groundwork for the gradual acceptance of Darwin's theory of evolution. The prevailing concepts of the more purely biological aspects of paleontology in 1850 were developed largely from the con- tributions of the above-mentioned investigators and thus were laid the founda- tions for a rapidly expanding science of paleontology. Historical Geology Before 1850 The prevailing ideas concerning stratigraphy at the opening of the nine- teenth century largely resulted from the influence of the earlier teaching of A. G. Werner in Germany. Contemporary publications were largely of a de- scriptive nature, with emphasis on places of occurrence, thickness of layers, and mineralogical composition of the rock. The importance of the use of fossils in determining the age of the strata had not been considered. The science of stratigraphy was established in England early in the nineteenth century as the result of detailed field studies by William Smith. Without formal training 692 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Smith devoted many years to the tracing of surface outcrops of different layers of slightly tilted strata across a part of England and collected and kept separate the fossils from each stratum. He recognized that each stratigraphic unit was characterized by a set of genera and species and that these faunas differed one from the other in the succession of beds from the base to top. He found that a single stratum when followed for long distances along the surface contained the same assemblage of species and that any particular isolated layer of rock could be identified by the fossils within it as belonging to a certain bed within a suc- cession of strata. The trends of each stratigraphic member were drawn in color on maps with sections showing the sequence of layers as they passed beneath the surface. These maps, published with an explanatory text (W. Smith, 1815), constitute the first large areal geologic map. It served as the foundation of a new epoch for the presentation of the results of field work and stratigraphic research to the scientific public, and opened a wealth of accurate information to the geologists of the Continent who were concerned with unraveling the geo- logic history of Mesozoic rocks. Conybeare and Phillips (1822), following the principles set forth by Smith, published the results of their investigations in England and Wales and classi- fied strata ranging in age from middle Paleozoic to Eecent. The terms Eocene, Miocene, and Pliocene later introduced by Charles Lyell were not employed but rocks corresponding to these ages were recognized and referred to according to their mineral composition. The Cretaceous was divided into upper and lower units and the Jurassic into the Oolitic system and the Lias, the former being subdivided into upper, middle, and lower Oolite. The New Red Sandstone be- neath the Oolitic was placed in the lower Mesozoic and above a fourfold division of the Carboniferous, the upper part of which was considered as equivalent to the Zechstein of Germany. The Old Red Sandstone was included within the Carboniferous until later assigned to the Devonian by Sedgwick and Murchison. The lower Mesozoic in Germany was studied in detail by various workers and divided into three members with the Bunter sandstone at the base, the ]\Iuschel- kalk limestone in the middle, and the Keuper at the top; in 1834 the name Tri- assic was applied to the group by Alberti. The absence of the marine middle member in Great Britain was early recognized by British geologists. In England, below the Old Red Standstone and above the granites and crystalline schists are a series of slates, limestones, and indurated sandstones which have suffered intense deformation. These rocks were studied by Sedgwick and Murchison and ultimately classified as the Cambrian and Silurian systems. The scarcity of fossils and the complexity of the stratigraphy rendered it im- possible at first to establish the boundary betw^een the two systems, with Murchi- son and Sedgwick becoming bitter enemies over the Cambrian. Murchison later (1839) included the entire assemblage of strata in the Upper and Lower Silu- rian and considered Sedgwick's Cambrian as a part of the Lower Silurian. Ulti- mately, however, Sedgwick's Cambrian was shown to be distinct from the Silurian. In 1879 the Lower Silurian rocks were studied by Lapworth and placed in a new system which he named the Ordovician. Meanwhile the Continental geologists examined the middle Paleozoic rocks in Germany, Belgium, Austria, and Scandinavia and attempted to make correlations with the succession of strata established in Great Britain bv Sedgwick and Murchison. Among the WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 593 more important of these investigations were those of C. F. Roemer (1844) on the faunas and stratigraphy of the Silurian and Devonian rocks of the Rhineland. The classification of the less deformed upper Paleozoic rocks in Great Britain into Old Red Sandstone, Carboniferous Limestone, Millstone, Grit, and Shales, and Coal Measures earlier in the century had been established by Conybeare and Phillips and named the Carboniferous system. The geographical extension of these limestones into Belgium afforded an opportunity for an intensive study of the faunas by de Koninck in 1844. The occurrence of similar limestones in the eastern Alps was noted by von Buch in 1824. The uppermost Paleozoic rocks in Germany were termed the Zechstein and consist of a succession of shales, sandstones, and conglomerates which pass upwards into limestone, dolomite, and marl. These rocks which underlie the Triassic were investigated early in the nineteenth century and the Zechstein group was considered as equivalent in age to the Magnesian Limestone of England. The rocks of late Paleozoic age in the Ural Mountains of Russia had been studied by several geologists and the result- ing published maps drew attention to this area as worthy of special investiga- tion. Accordingly, Murchison, who already had examined many areas in the Alps and other parts of Europe, was requested by the Russian government to make a geological study of the Province of Perm. He was accompanied by the French geologist, de Verneuil, and the Russian Count, von Keyserling. The results of this investigation were published in 1845 in their monograph. The Geology of Russia in Europe and the Ural Mountains. The rocks which overlie the coal-bearing beds were called the Permian system. The marked variations in the lithologic character of the faunal facies of the Cretaceous rocks in different parts of Europe in contrast to the greater uni- formity prevailing among the Jurassic resulted in the development of several distinct forms of classification and nomenclature. Numerous important contri- butions were published before the middle of the nineteenth century and the succession of formations in each country was fairly well established. By 1850 the broad outlines of the stratigraphical and faunal classification of the Creta- ceous deposits were somewhat similar to those in use at the present time and served as a foundation for more detailed research in other parts of the world from 1850 to 1950. In England the sedimentary rocks lying between the basal Tertiary and Upper Jurassic were classified by William Smith in downward sequence as the White Chalk, Gray Chalk, Greensand, and Micaceous Clay. Certain dark claj'-s occurring locally beneath the Greensand were called the Blue Clay but were later designated the Gault. Later investigations by Fitton in eastern Sussex showed that a portion of the lower Greensand passed from a marine to fluvia- tile facies. This was termed the Wealden formation. These deposits contain a rich flora and the remains of fossil reptiles. The name Speeton clay was given to beds in Yorkshire equivalent to the lower Greensand. The Upper Cretaceous or White Chalk was recognized as extending eastward into Belgium, France, Den- mark, North Germany, and Poland. The Cretaceous stratigraphic and faunal units of Great Britain are widely represented in France but the complete sequence of beds is not everywhere pres- ent. There is a prevailing similarity of the upper White Chalk and marls but the middle and lower members often present marked differences in litliology. Early 694 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES in the nineteenth century Lower Cretaceous shales, limestones, and marls had been recognized as resting on uppermost Jurassic deposits in the Jura Alps of eastern France and in 1835 these rocks and their fauna were described and desig- nated as Neocomian. The Cretaceous rocks and faunas in particular areas of France were studied by Leymerie, d'Archiac, and others and compared and cor- related with the divisions established in England, Germany, and Switzerland. During the first half of the nineteenth century, the Cretaceous deposits of Germany were studied by F. A. Roemer and Hans Geinitz. The former in 1841 published his great work on the Cretaceous of North Germany, with a descrip- tion of the faunas and a classification of the rocks. At approximately the same time Geinitz described the Cretaceous faunas and rocks of Saxony and Bohemia. By the middle of the century the first volumes of d'Orbigny's PaUontologie frangaise had appeared. In this work the French Cretaceous was divided into seven stages designated as Neocomian, Urgonien, Aptien, Albien, Gault, Ceno- manien, Upper Greensand, Turonien, and Senonien. Additional investigations by Geinitz and Beyrich extended the German classification to the Cretaceous deposits farther east in Hungary and north into the Baltic region. The results of early investigations of the richly fossiliferous Tertiary deposits of the Paris Basin led to a preliminary classification of the strata and a knowl- edge of the stratigraphic relationships of the invertebrate faunas occurring in the beds which later were to be designated as Eocene and Oligocene. An impor- tant contribution by Cuvier and Brongniart was published in 1808 and reprinted in 1811, in which the faunas were listed and the rocks described. Accompanying preliminary geologic maps and stratigraphic sections revealed the relatively simple structural features of the strata and some information concerning chang- ing lithologic facies. The recognition of lower beds of plastic clay, followed by sandstones and limestones forming a group characterized by great numbers of Nummulites, led to the introduction of the term Nummulitic Series for strata which later were to be classified as Eocene. Above was a second group, which in upward succession consisted of gypsum, fresh water marls, and clays, passing upward into marine limestones and sandstones alternating with fluviatile and lacustrine beds. From such evidence Cuvier and Brongniart concluded that at the close of the Cretaceous, the area of the Paris Basin was only slightly above sea level and that it possessed an irregular surface formed by erosion of the Up- per Chalk and that it was traversed by several streams. A local, differential, slight downwarping of the surface at the close of the Cretaceous permitted ma- rine waters to transgress slowly into the river valleys and ultimately over the intervening land areas, thus permitting the deposition of fluviatile and lagunal deposits and the plastic clay. Intermittent advances and retreats of the sea were regarded as being responsible for the alternations of continental and marine beds and variations in sedimentary facies. The Tertiary of the Belgian Basin was laid down in the northeast extension of the Paris Basin. One of the most important studies on these deposits was by A. Dumont (1849) in which the entire Belgian Tertiary succession was classi- fied stratigraphically upward as the Heersien, Landenien, Ypresien, Paniselien, Bruxellien, Laekien, Tongrien, Rupelien, Bolderien, Diestien, and Scaldisien. Among the more important contributions made to the Tertiary stratigraphy and paleontology of Italy was a work published in 1814 by Giovanni Brocchi V/EAVEk: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 595 on the Subappenine fossils. These were described and illustrated and a record was made of their exact occurrence within the different strata from which they were collected. Attention was called to the biological similarities of some of the species to those now living in the Mediterranean and also to certain forms from the Paris Basin. Other investigations by Omalius d'Halloy in Belgium and other parts of southern Europe and Germany presented evidence for future corre- lation of the Tertiary deposits from different parts of the continent. Among the earlier publications dealing with fossils and rocks of Tertiary age was the work by Conybeare and Phillips in 1822 in which the sedimentary deposits in Great Britain above the Cretaceous were classified in ascending order as the Plastic Clay, London Clay, Freshwater Beds, and the Upper Marine For- mation consisting of the Bagshot Sands and the Crag. This work, along with numerous others including the description of fossil mollusks by the Sowerbys, afforded material for a comparison of the British Tertiary fossils with those described by Deshayes in 1824-1837 from the Paris Basin, Italy, and other parts of Europe. Deshayes recognized an increase in the proportion of living species among the successive faunas from earliest to latest Tertiary. During this time Charles Lyell, who had traveled extensively in France, Italy, and other parts of Europe, studied the Tertiary outcrops and also the fossil collections that had accumulated in the museums and universities. Working independently, H. G. Bronn made detailed investigations of the Tertiary rocks and faunas of Italy and also observed the increase in the percentage of living species and genera in the younger beds. Lyell, after considerable association with Deshayes in Paris, correlated the deposits of France with those in the south of England and, influ- enced by the increasing percentage of living species from the base to top of the Tertiary, established new terms for the major divisions. In the third volume of his Principles of Geology published in 1833 he introduced the names Eocene, Mio- cene, Pliocene, and Recent. In 1846 the name Pleistocene was introduced by Forbes to include the uppermost Pliocene of Lyell and deposits of glacial origin. After 1850 the additional divisions of Paleocene and Oligocene were added by other workers. Geological and paleontological research in eastern North America during the first half of the nineteenth century was carried on largely under the auspices of the newly established state geological surveys and by individuals associated with a few universities and academies of sciences. Ebenezer Emmons, after a study of the rocks in New York State, classified them in upward succession as tlie Crystalline complex, the Taconic system of greatly disturbed beds, the New York system, and the Red Beds. The New York sj^stem consisted of nearly horizontal beds of sandstones, shales, and limestones, which he subdivided into the Champlain, Ontario, Ileldeberg, and Erie groups. A report published by James Hall of the New York Survey in 1843 subdivided the New York system into twenty-nine groups. The Paleozoic rocks of Pennsylvania were investigated by H. D. and W. B. Rogers, who in 1843 described their composition, lithology, thickness, and geologic structure. The coal in the upper beds was considered as having been formed from peat bogs occurring on the surface of an extensive plain which from time to time suddenly passed slightly below and above the level of the sea. The Rogers recognized the folded character of the Appalachian Range and considered its deformation to have taken place after the deposition 696 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES of the coal measures. In 1847 James D. Dana attributed the folding to horizon- tal compression of the strata as the result of the slow cooling and contraction of the earth's nucleus. The foundations of Tertiary stratigraphy and invertebrate paleontology in North America were established by the investigations of T. A. Conrad with a series of contributions commencing in 1832 and continuing to and beyond 1850. The more important of his earlier papers included descriptions and illustrations of the Eocene shells of Alabama, the medial Tertiary fossils of the Atlantic Coast, and studies of fossils collected by W. P. Blake in California, as well as those obtained by the Wilkes Exploring Expedition near the mouth of the Columbia River at Astoria, Oregon. Many of these contributions were pub- lished in the American Journal of Science, the Journal of Conchology, and the Proceedings of the PMladelpJiia Academy of Sciences. The visit of Sir Charles Lyell to America in 1844 rendered possible a comparison of the Tertiary de- posits with those of Europe. Reseakch and Publication Facilities Prior to 1850 The earlier contributions to paleontology and historical geology in Europe were largely by men who were sponsored and financed by those in control of state governments or by financially independent individuals interested in scien- tific work. After the private publication of William Smith's geological map of a part of England the value of geological work by the state became apparent. The Geological Survey of the United Kingdom was founded in 1835 under the direction of de La Beche and about the same time preparations were begun for a geological map of France. In North America the State Geological Survey of North Carolina was established in 1823, other states following this example, South Carolina in 1824, and Tennessee in 1831, while the Canadian Geological Survey was founded in 1841. The universities in Europe have always taken an impor- tant part in advancing the sciences of paleontology and historical geology and in training future investigators. Collections of fossils, together with library fa- cilities, rendered possible the comparison of materials gathered from various parts of the world. Toward the middle of the nineteenth century geologists and paleontologists established professional organizations which provided scientific meetings, where the results of investigations could be presented and discussed, as well as facilities for publication. The Geological Society of London was or- ganized in 1807 and the Palaeontographical Society of London in 1847. Impor- tant monographs on both invertebrate and vertebrate fossils began to appear in 1847 in the German publication Palaeontographica. The Geological Society of Germany was founded in 1849 and in it have appeared many important contri- butions to the geology and paleontology of Europe and other parts of the world. Invertebrate Paleontology 1850-1950 At the beginning of the second half of the nineteenth century the theory of the immutability of species, together with the idea of special creation, had been strongly challenged by an increasing number of scientific investigators. Many important papers and monographs had been published with descriptions and il- WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 697 lustrations of genera and species. The morphology of the hard parts of different groups of invertebrate organisms was now well known; also the facts connected with the occurrence of fossils in successive layers of sedimentary rocks. Infor- mation concerning their geographic distribution and their relation to changing lithologic facies caused many investigators to accept partially the new concept of mutability of species. Those who now did so no longer considered a species as confined to a single formation or interval of geologic time but held that cer- tain species of a fauna might survive and continue to exist in the next succeed- ing set of strata. These concepts profoundly modified the ideas previously held by scientific men in the fields of paleontology, geology, and natural history. Speculative concepts of the principles of evolution had been expressed before but were not generally supported by scientific facts. Paleontological research was now endeavoring to inquire into the nature of evolutionary processes based on a growing knowledge of the succession of faunas through geologic time. Charles Darwin began his investigation of the origin of species during this inter- val of change in scientific thought and in 1859 his monumental work on The Origin of Species by Means of Natural Selection appeared. The ideas expressed in this work appealed strongly to H. G. Bronn, who had already considered that transitional genera and species in a fauna of a given time interval might extend across the gap into the next succeeding interval. He translated Darwin's work into German and it immediately influenced the thought of those scientists on the Continent who were concerned with paleontological investigations. Darwin, in attempting to account for the method of action in the theory of evolution, placed emphasis on those factors involved in environment, including variations in climate, temperature, accessibility of food supplies, and means of protection from other organisms living in the same environment. He considered also how these factors singly or jointly might react favorably or unfavorably on each in- dividual, depending on how each particular feature of the anatomy might re- spond. Morphological characters of a varietal nature were considered to influ- fiuence the individual's response to the varying stimuli of the environment; thus those individuals with the most favorable variations would survive. This con- cept was designated as the law of natural selection. The principles of sexual selection were also included. Before leaving for North America in 1847 Louis Agassiz had published the results of ten years of research on fossil fishes. Emphasis was laid on genealogy, including progressive morphological changes of certain parts of the skeleton as observed in different species and genera in passing from older to younger time intervals. Agassiz showed that information from the phylogenetic history could be of great value to geology for the systematic classification of sedimentary de- posits. He also pointed out the correspondence of certain changes in the embry- onic development of an individual to those exhibited in the phylogeny during geologic time. During the second half of the nineteenth century, as a lecturer in Harvard University, he inspired many of his students with the spirit of re- search and the inclination to search further for more detailed factors involved in the process of evolution as shown in fossil organisms. The detailed morphological study necessary to yield sufficient data for ex- panding these principles of evolution of organic life during past geologic epochs required that the investigator devote special attention to the fossils of a single 698 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES phylum or subdivision of it. Accordingly the more important advances made during the past one hundred years have been centered on the study of special groups, with particular attention to their geologic history. Many monographs dealing strictly with the biologic aspects of faunas were, and still are being, published but some of the more important, while contributing to the problems of evolution, lay special emphasis on stratigraphic problems. The important advances made in the science of invertebrate paleontology during the past one hundred years may best be outlined by the consideration of each invertebrate phylum separately. Protozoa: The foraminifera, which in most cases have chambered tests, in 1798 were included by Cuvier in the Mollusca but later, when studied micro- scopically by Dujardin in 1835, were found to contain protoplasm which was free to circulate throughout the chambers, thus establishing their unicellular structure. During the second quarter of the nineteenth century d'Orbigny de- voted much time to a study of the Foraminifera and in 1826 made this group of organisms a special order, which he included in the class Cephalopoda. The order was divided into 52 genera and over 500 species based largely on the mor- phological differences of the wall and the shape of the test. He later accepted the interpretation of Dujardin that the Foraminifera were protozoans and in 1839 revised the earlier classification so as to consist of 6 orders and 64 genera. His final paper in 1852 grouped the genera into 7 orders, in which the growth and arrangement of chambers were of fundamental importance for classifica- tion. He still adhered to the concept of immutalibity of species. The foraminiferal investigations initiated by d'Orbigny were continued by numerous paleontologists, and many classifications were devised, based on dif- ferent criteria as the description of new genera and species appeared in a rapidly expanding literature. M. S. Schultze in 1854 proposed that all of the foraminiferal genera be included in a single order Testacea, which he separated from the rest of the unicellular organisms of the phylum Protozoa. The species were included under 10 families and 112 genera, all classified according to the shape of the test and arrangement of chambers. AVilliamson, who devoted spe- cial attention to a detailed study of the structure of the hard parts, concluded that the great differences shown in the structural characters rendered difficult any exact classification. He regarded these differences as due largely to varia- tion and arranged into groups those genera which appeared to be closely related and grading into each other. A classification founded on the number, structure, and arrangement of chambers of the test, as well as the perforate or imperforate character of the wall, was proposed by A. E. Keuss in 1861. In addition, differ- ences in the chemical and mineralogical composition were included. All of the species were grouped into 21 families and 109 genera. H. B, Brady (1884), who studied the Eecent foraminifera obtained by the "Challenger" Expedition, constructed a classification consisting of 10 families and 153 genera based largely on the texture, shape and structure of the test, aperture, and number and arrangement of the chambers. He was not in agree- ment with Reuss, who had included all arenaceous foraminifera in the imper- forate groups, because some of the genera possess mural pores. The Astrorhizi- dae were considered by Neumayr (1887) as a primitive group and as the ances- tors of the other foraminiferal families already established by Brady. On the WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 699 basis of this interpretation Neumayr built up a phylogenetic classification based in part on the' stratigraphic range of species and genera. Ludwick Rhumbler (1899) considered that the evolutionary development of foraminifera during geological time involved morphological changes which would produce a strengthening of the test and that the uniserial forms would be more easily injured or destroyed than the coiled types. Accordingly, he regarded the former as ancestral to the more complexly coiled forms, a conclusion which is not entirely in agreement with the information concerning the occurrences in the fossil record. The evolution of the foraminifera was discussed in an important publication by Henri Douville (1907). In this he considered that the composition of the test was dependent to a large extent on the differences of environment under which the animal lived, that the calcareous forms were descended from the are- naceous, and that those with uniserial tests represented a more advanced state of development than the coiled forms. At present the arenaceous types are not considered ancestral to the calcareous. The classifications presented by Cushman from 1927 to 1940 raised the number of families to 49, which Cushman believed were derived largely from coiled, nonseptate genera; he also thought that the calcareous group had arenaceous ancestors. His classification accepts many of the principles advocated by Douville. Galloway in 1933 criticized Cushman's classification, asserting that the latter had established too many families and subfamilies and that many of the genera had been placed in wrong families and in incorrect relationships between families. Galloway's classification and ar- rangement of families were based on his interpretation of the phylogenetic char- acter of the different groups and determined from a consideration of comparative morphology, the stratigraphic range of the different assemblages of genera, and the biogenetic law. He considered the foraminifera as consisting of two broad groups, one of which evolved from the Allogromiidae, the other from Endothyra. The former constitutes a group of foraminifera with a single chamber and, be- cause of its chitinous wall, is rarely preserved as a fossil. However, he consid- ered it ancestral to the nonseptate calcareous and arenaceous forms, including the Miliolidae. The most recent classification is by M. F. Glaessner, published in 1945. The order Foraminifera is divided into 7 superfamilies, 37 families, and 38 subfami- lies. Glaessner concludes: 1. The non-septate forms are more primitive than the septate. 2. The higher, or septate, spirally coiled arenaceous foraminifera form a well-defined group. 3. The Fusulinidae are derived directly from Endothyridae. 4. The different lines of the porcellaneous foraminifera have a common origin in a coiled non-septate form. 5. The Polymorphinidae are derived from Legenidae but there is no clear evidence concerning the origin of this family. 6. The Cassidulinidae and Ellipsoidinidae (Pleurostomellidae) are related to the Buliminidae which can be traced back to a trochospiral ancestral form. Most of the other smaller calcareous perforate foraminifera are clearly derived from rotaloid (trochospiral) ancestors. 7. Most of the larger calcareous perforate foraminifera including Siderolites, Orbi- toides, Lepidocyclina, Miogypsina, and probably the nummulites, developed from a number of different but closely inter-related small rotaloid (trochospiral) ancestors. 700 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES The ever-increasing demand for gas and petroleum products over large areas of the world intensified stratigraphic investigations and paleontologic research. Foraminifera began to play an important role in the determination of the geo- logic area of strata associated with the occurrence of oil. Thick deposits of shale with a wide areal geographic distribution often are nearly barren of diagnostic metazoan invertebrate fossils but are rich in foraminifera and other microscopic organisms. Because of their small size, well preserved faunas consisting of a vast number of individuals may be obtained from small samples of rock such as drill cores. The growing demand for scientifically trained investigators in micro- paleontology soon led to the introduction of special courses in many universities where opportunity was available for instruction in this field. The larger oil companies began to add micropaleontologists to their geological staffs and some companies established special departments with well-equipped laboratories for handling this kind of work. Many important contributions to foraminiferal re- search have resulted from investigations carried on by those associated with the oil industry. Not only the universities of North America but those in other parts of the world as well have contributed to the description of new species and genera and their stratigraphic relationships. Increasing attention is being devoted to this kind of research by national and state surveys with the publica- tion of many important papers and monographs concerned with foraminiferal faunas. Such investigations have been augmented by scientific and professional societies and academies throughout the world. Conspicuous among those in North Ameria are the Cushman Laboratory for Foraminiferal Research, the Paleontological Research Association and the Paleontological Society, and the Society of Economic Paleontologists and Mineralogists. As the result of investigations during the past one hundred years there have been described about 4,000 species of Foraminifera distributed through nearly 300 genera. The practical application of micropaleontology has involved the use of species in local basins for deciphering the succession of strata as exposed at the surface and in the recognition of such beds under a deep covering of younger deposits as revealed in the cores obtained by drilling. Only a part of this de- tailed information has been available to those paleontologists concerned with the more purely scientific aspects of investigation, including the regional distribu- tion, stratigraphic succession over wide areas, and ecological relationships. How- ever, an increasing record of statistical data in recent years has made possible the publication of important papers dealing with an understanding of the ecology of assemblages, the influence of environmental changes in the composi- tion of faunas from the study of living species, variation in species, and world- wide succession of faunal zones. From such research, modifications are being made in phylogenetic classification and in the evolution of different groups of foraminifera. Porifera: In the early part of the nineteenth century there was uncertainty whether sponges were animals or plants. The group was investigated by Robert Grant of England, who in 1825 ascertained their affinity with the animal king- dom. Seventy-five species were described by Goldfuss (with G. Munster) in his Petrefacta Germaniae published in 1826, but the morphology of the group was still little known. Fossil sponges from the Upper Cretaceous of England were studied by Toulmin Smith in 1847-1848 with special attention to the structure WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 701 of Ventriculites, which he considered related to the Bryozoa. In 1852 d'Orbigny classified the sponges as Amorphozoaires, dividing them into two orders: (1) Amphozoaires a squelette corne and (2) Amphozoaires a squelette testace. The first order included living forms and the fossils belonging to the genus Cliona; the second order was divided into five families. The finer morphological charac- ters of the skeleton were not considered, and his classification was based largely on external features. E. de Fromentel (1859) in his introduction to the study of fossil sponges — both living and fossil — considered the canal system, pores, osculum, and tubules for classification purposes. Sponges from the Jura Moun- tains were investigated by Etallon in 1859 and 1861, with special attention to the spicules, canal system, and the outer form, all of which he considered of im- portance for classification. Ferdinand Roemer (1860) studied the Silurian sponges of western Tennessee and defined the genera Astylospongia, Palaeoma- non, and Astraeospongia. In 1864 F. A. Roemer published a monograph on the sponges of the North German Cretaceous, with the description of many species and an excellent account of the structure of the skeleton of the Hexac- tinellids and Lithistida. Sponges from the Miocene rocks of Oran were described in 1872 by A. Pomel, who developed a classification consisting of two broad groups, (1) Camptonspongiae and (2) Petrospongiae. The first, with two orders, was characterized by isolated spicules; in the second, with 239 genera, the spicules were arranged in a united framework. Wyville Thompson, who studied the sponges collected by the "Challenger" Expedition, was the first to point out a similarity of structure in the fossil Ventriculites to that in living siliceous sponges. In the last quarter of the nineteenth century considerable advance was made in the study of sponges as the result of the examination of thin sections under the microscope. By this method SoUas in 1877 compared several genera from the English Chalk with living hexactinellids and monactinellids. Similar studies by von Zittel (1877-1878) led him to consider that all sponges, both fos- sil and living, should be included in a single classification. Hinde, in the light of increasing knowledge of the morphology, presented a classification very close to that now in general use. It included the four orders Myxospongia, Cerato- spongia, Silicispongia, and Calcispongia. A discussion of the advances made in the investigation of fossil and Recent sponges, along with a more advanced scheme of classification was given in a monograph by Rauff in 1893-1894. This grouping is used in the translation by Eastman (1913) of von Zittel's Inverte- hrate Paleontology, in which the sponges are placed in the phylum Coelenterata along with the corals but are included in the subphylum Porifera and in the class Spongiae. The four subclasses Myxospongiae, Ceratospongiae, Silicispongiae, and Calcispongiae are still recognized but the first two, because of their lack of imperishable hard parts, are not included in the textbook. The Silicispongiae are divided into four orders, the last two of which include the majority of fossil sponges. The Calcispongiae include two orders, Pharetrones and Sycones. The classification is based largely on differences in the character of spicules, whether single or united into a framework, thickness of walls, character of pores and tubes, and the osculum. Monographs dealing with special groups of sponges have contributed greatly to the advances made during the past sixty years. Among the more important of t;hese is the work on the Dictyospongiae-Paleozoic Reticulate Sponges, which was 702 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES published by James Hall and J. M. Clarke (1898). These sponges are particu- larly abundant in the Devonian of New York State. In 1842 T. A. Conrad de- scribed and named Hydnoceras tuberosum and considered it a cephalopod. Lard- ner Vanuxem described TJphantaenia chemungensis as a marine plant. Another closely allied form was called Dictyophyton, revealing these authors' concept of its place in the classification. Many diverse forms were described before 1880 but the conditions of preservation made recognition of their real nature difficult. The internal and external casts of their bodies left distinct impressions of their spicular network. Similar fossils from Crawfordsville, Indiana, yielded more definite evidence and led to a recognition of their relationship to the living reti- culate silicious sponges represented by Euplectella aspergiUum. A group of Cambrian sponges of the class Pleospongia were investigated in great detail by Vladimir J. Okulitch (1943). His classification is based on their morphology, affinities, and distribution. The first representatives of this class were described in 1861 by E. Billings of Canada as Archaeocyathus from the Lower Cambrian of Labrador. Okulitch 's classification consists of 5 subclasses, 11 orders, 20 families, and 89 genera. Previous to the appearance of this work the Pleospongia were variously grouped with the Foraminifera, Sponges, or Corals. Okulitch concludes that they represent a separate class of the Porifera which became extinct in the Upper Cambrian, that there are no links between the Silicispongia and Calcispongia, and that they should be regarded "as inde- pendent branches having a common ancestor in the Pre-Cambrian." Coelenterata: The Coelenterata is a phylum more advanced than the sponges and consists of a large number of extinct and living species. They are repre- sented in the fossil record from the Cambrian to the Eecent. At present they are usually divided into the following groups: Ilydrozoa, Stromatoporoidea, Grap- tozoa, Scyphozoa, and Anthozoa. Many genera and species of corals, both liv- ing and fossil, were described in the first decades of the nineteenth century but the organization of their morphological features was not well known. Among the earlier authors were Lamarck, Ehrenberg, and Goldfuss. Thorough investi- gations of living corals were made by ]\Iilne-Edwards and Haime. These were followed by special studies of particular groups of fossil forms, with special at- tention to the morphology and structure of the polyps and the occurrence and distribution of fossil faunas through geologic time, culminating in the Histoire natureUe des coraUaires (1857-1860). The classification of Milne-Edwards and Haime is based on the differences in the septa and the methods by which new ones are produced. The subdivisions are based on the number of tentacles. During the middle of the nineteenth century many papers, largely of a de- scriptive nature, were published, among the more important of which were those by Reuss, Fromentel, Hall, and Duncan. Differences in the method by means of which new septa originated in the Paleozoic Tetracoralla as compared to those of the Hexacoralla and Octocoralla were described by Kunth in 1869-1870. Thin sections made from the calcareous skeletons were subjected to microscopic analysis. A similar method of study was used also by Pratz and Koch with ac- companying illustrations in their published work. In 1896 Maria Ogilvie investi- gated fossil and living anthozoans and presented her interpretations of the phy- logenetic relationships of the Tetracoralla and Hexacoralla. Moseley (1877) in a study of MiUepora showed the relation of Stromatoporans to certain Hydrozoa SNEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 703 which earlier had been considered Bryozoa. Nicholson's (1886-1892) important monograph on all known Stromatoporoids was published in the 'nineties and has been followed, with many modifications, by later authors. The fossil ' Stromatoporoidea consist of large calcareous masses with great variation in structure and shape. Their structure, based largely on fossils from western Germany, has been investigated by M. Heinrich (1914), who divided these organisms into two groups dependent on the massive or nonmassive character of of the fibers. Those forms having a regular rectilinear arrangement of the fibers were placed in the massive group and those with an irregular vermiculate ar- rangement of the hollow-fibered skeletal structures in the nonmassive groups. The principal characters used in both groups for distinguishing genera include the amount of regularity and varying pattern of the skeletal mesh. An important contribution to the Anthozoa was published in 1900 by T. W. Vaughan, with special emphasis on the general character and bathymetric distri- bution of Eocene and Oligocene genera and species in North America. He consid- ered that the classification of corals was in a very unsatisfactory condition, that "no classification that will stand the test of thorough criticism has yet been pro- posed . . . [and that] past classifications were based on some particular features of the skeleton without reference to the whole structure and history of the organ- ism." He pointed out that Duncan had based his grouping on a combination of general skeletal features and mode of growth but had not searched for those char- acters which were of phylogenetic importance. The accumulated evidence that the Tetracoralla were confined to the Paleozoic and the Hexacoralla to later geologic time led P. E. Raymond to consider that the cooler climate of Permian time had brought about the extinction of that group largely because they had become specialized as lime-secreting organisms in rela- tively high-temperature seas. He also proposed the idea that the Hexacoralla origi- nated from an "Edivardsia-\ike actinian" of the Paleozoic and became a lime- secreting organism as the temperatures lowered at the close of the Permian. Robin- son suggested that certain forms classed as Tetracoralla occurring in the Meso- zoic were probably Hexacoralla and that a number of Paleozoic genera referred to the Hexacoralla were Tetracoralla. This pointed to a sharp distinction in the time relations of these two groups. The paleozoic Cyathaxonia was sug- gested as a specialized organism which might have evolved into Mesozoic types, in which through assumption of an upright position the columella would change from an excentric to a central location. The graptolites, which are abundant as colonies in black shales of the early Paleozoic where they are compressed in the rock layers like fossil leaves, occur as individual polyps attached to a central axis. They are important for distin- guishing the geologic age of Ordovician and Silurian rocks and were placed by many early authors, including Nicholson and Lapworth, with the Hydrozoans. However, Neumayr held that they could not be placed with any of the known classes of animals. In 1931 E. 0. Ulrich and R. Ruedemann, who had investigated the graptolites for many years, discussed the question whether they should be classed wdth the Hydrozoans or Bryozoans. Their objections to placing them with the Hydrozoans were based on the ground that the Hydrozoa contain no structures in common with graptolites, that they exhibit a different type of symmetry, and that the graptolites include "a considerable number of unnaturally associated 704 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES fossil types." Ulrich and Ruedemann considered that the graptolites were more closely related to the Bryozoans because of the character of the sicula, the mode of budding from the sicula, the bilateral symmetrical thecae, and the similar habitus of the two groups. Recently R. Kozlowski (1948), in studying excellently preserved material from Lower Ordovician cherts in Poland, has concluded that the graptolites are closely related to the living Pterobranch Rhahdopleura of the phylum Hemichordata. The fossil Medusae, which usually possess only a slight resemblance to living medusae, have been described in an important monograph by C. D. Walcott (1898). During the second half of the nineteenth century many species were figured and discussed by Beyrich, Haeckel, Ammon, and Nothorst. E chinodermata : The accumulated information of 1850 concerning the mor- phology and classification of the phylum E chinodermata had resulted from in- vestigations carried on in both Europe and North America. The Crinoidea were established as an independent group by J. S. Miller in 1821, the Blastoidea by Fleming in 1828 and the Cystoidea by von Buch in 1845. In 1848 Leuckart combined these under the class name Pelmatazoa and placed it along with other echinodern groups in the phylum Echinodermata. Important monographs by Goldfuss and Munster in 1826, Johannes Miiller in 1841, Vaughn Thompson in 1836, Edward Forbes in 1848, and d'Orbigny in 1840-1860 emphasized the mor- phological details of genera and families along with the description and illus- tration of new species with attempts at broad classifications. From 1850 to 1950 a very extensive literature accumulated concerning the morphology and classi- fication of echinoderms and the relationships of genera to their stratigraphic succession. The earliest classifications of the Pelmatazoa were not founded on morpho- logical principles, and many greatly differing forms were combined and placed in the same group. The Blastoids and Cystoids were placed by those immediately following J. S. Miller as a subordinate group under the Crinoids. Miller had divided the Crinoids into four groups, largely on the number and arrangement of the plates in the dorsal cup. Most investigators following Johannes Miiller considered that all Paleozoic forms were distinct from later ones and it was not until the publication of Carpenter's work in the "Challenger" reports in 1884 on the stalked Crinoids that the morphological relations between the Mesozoic and Paleozoic forms became known. He held that the Paleozoic Crinoids dif- fered from those of later age in the character of their irregular symmetry. Many new species and genera of Crinoids and Cystoids from Bohemia were de- scribed by J. Barrande between 1877 and 1899. In 1845 von Buch gave the Cystoidea equal rank with the Blastoidea and Crinoidea. Ferdinand Roemer in 1855 published an important memoir on the Cystoidea and Blastoidea and divided the former into three groups. Neumayr considered the Cystoidea, Blas- toidea, and Crinoidea independent classes and believed that the two last were derived from the Cystoidea. During the fourth quarter of the nineteenth century the Pelmatazoa were the subject of intensive investigation by North American paleontologists. Among the more noteworthy were F. B. Meek, A. H. Worthen, E. Billings, James Hall, Charles Wachsmuth, and Frank Springer. Under the auspices of the Geological Survey of Canada Billings, in 1869 and 1870, published contributions on the WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 705 structure of these forms, with special attention to the position of the mouth in relation to the ambulacral system. He concluded that the pores in the rhombs of the Cystoidea were respiratory organs and the homologues of the tubular apparatus which underlies the ambulacra of the Blastoidea. He pointed out that Eocystites was the most primitive genus of the Cystoids, with an indefinite num- ber of plates without any radial arrangement. He showed that the hydrospires of the Blastoidea were connected in pairs and had direct communication with the pinnulae. He thought that the mouth and anus combined represented the opening in the disk of Paleozoic Crinoids and that the grooves which pass from the center of the disk at the inner floor were conected with the ambulacral sys- tem and communicated through the arm openings with the arm grooves but did not enter the tegminal aperture. He considered that the food entered the body through the arm openings and was carried underneath the tegmen to a common oral center. Meek and AVorthen added to these morphological studies, with at- tention to the ventral surface of the calyx, the plates, mouth, and anal openings. In North America little attention had been given to the Crinoids until 1858 but by 1897 over 1,400 species had been described. Until the publication of the "Challenger" reports studies were confined largely to the abactinal side of the calyx and no attempt was made to homologize the plates of the tegmen of the different groups. The excellently preserved and abundant faunas of the late Paleozoic in the Mississippi Valley drew the attention of Wachsmuth and Springer, who, during the last quarter of the nineteenth and first quarter of the twentieth centuries, contributed many important papers and monographs on the Pelmatazoans. Among these was a three-volume work published in 1897 by the Museum of Comparative Zoology on The North American Crinoidea Ca- merata. Special consideration was given to morphological and phylogenetic characters, with an accompanying classification which is widely accepted at the present time. They point out "that the Crinoids were most intimately connected from the Silurian down to the present and only the Camerata, a highly special- ized type, became extinct at the close of the Paleozoic." All American species of the Camerata known at that time were described in this monograph. They noted that the Crinoids, Blastoids, and Cystoids differ from other Echinoderms in being at one stage of life provided with a stem for attachment to other objects, thus living on their abactinal side in contrast to the other groups. Other im- portant later contributions have been made by Jaekel on the phylogeny of the Pelmatazoa, by the revised textbooks of von Zittel, and by R. S. Bassler on the Edrioasteroidea. Although many genera and species of Echinoids were described during the first half of the nineteenth century, the scientific approach to the investigation of the morphology and phylogeny began with the contributions made by Agas- siz and Desor in 1840. These were followed during the next seventy-five years by the publications of Forbes, d'Orbigny, Wright, and Cotteau in Europe and by Hall, W. B. Clark, Twitchell, and Jackson in North America. The compre- hensive monograph by Jackson in 1912 on the phylogeny of the Echini presents the information and interpretations made by numerous authors both in Europe and America, together with his own study of the morphology, development, and comparative anatomy of this class founded on the young and adult of fossil and living forms. His contribution includes a revision of the Paleozoic Echini and 706 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES a systematic description of all known Paleozoic forms. He assumes that any scientific classification should be based on the totality of characters and not on single phases of the morphology. Emphasis is laid on the use of stages in de- velopment and a comparison of these with the characters of more or less closely associated types. He developed the idea of localized stages in development, the idea that, "throughout the life of the individual, stages may be found in definite parts that are comparable to the condition in the young and to adults of simpler types of the group." This principle was found applicable to such other groups as Crinoids, Corals, and Cephalopods and has been used by Hyatt, Beecher, Cushmann, Kuedemann, H. L. Clark, and many others. Jackson points out that senescence is well shown in Paleozoic Echini by the dropping out of columns of interambulacral plates at the dorsal portion of the test. He defines progressive types as "those which show in their development to maturity the addition of dif- ferential characters only ..." without their later disappearance and points out that most Paleozoic Echini are of this type. Regressive types are considered to be those which show specialized characters in later development but lose these before old age so that the adult is simpler than its ow^n young. Other characters discussed by Jackson are acceleration of development, parallelism, and variation. The more important factors used by Jackson in considering the comparative morphology of the Echini were the form of the test, the pentameral system, the structure of the skeleton, the ambulacra and interambulacra of the corona, the spines, peristome, and occular and genital plates. His classification of the Echi- noidea consists of 7 orders and 17 families. After a critical study of plate structure, Sven Loven in 1874 devised a nomen- clature for referring to the ambulacra in terms of the Roman numerals I to V and the interambulacra by the Arabic numerals 1 to 5. He established the method of determining the bilateral symmetry of the test by the presence of the madrejjorite or the periproct. Important contributions were made to the Mesozoic and Tertiary Echinoids in North America by W. B. Clark in 1893, W. B. Clark and M. W. Twitchell in 1915, W. S. W. Kew in 1920, and Grant and Hertlein in 1938. Cenozoic Echi- noids have been described by P. M. Duncan from Australia and by Duncan and Sladen from the Western Sind of India. The papers by J. Lambert and P. Thiery (1909-1925) are important for their taxonomic work on the Echinoidea. Fossil starfish are not extensively used for purposes of geological correlation, yet they occur in rocks from the Ordovician to the present. Many papers have been written describing genera and species, among the more important of which are those of Sladen and Spencer from 1890 to 1908 on the British fossil Asteroi- dea, F. Schondorf from 1907 to 1913 on the German Paleozoic forms, and a monograph by Charles Schuchert published in 1915 on the Paleozoic Stelleroi- dea. In this volume there are recognized 45 genera and 110 species. Fifty-one of the species are from North America, 53 from Europe, and 6 from the southern hemisphere. This paper is of special importance for its presentation of the skele- tal terminology of the Asteroidea. Brachiopoda : The class name Brachiopoda was proposed in 1802 by Cuvier. A memoir by von Buch in 1834 contained a classification based largely on the characters of the hinge area. The character of the brachial appendages, the septum, the muscular impressions, and other internal structures were used by WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY JQJ King (1846) in the construction of a classification in which the Brachiopods were subdivided into 3 orders, 16 families, and 49 genera. The important mono- graphs by Thomas Davidson published at intervals from 1851 to 1885 present an excellent analysis of the morphological characters of the hard parts of both fossil and living Brachiopods, along with a description of the soft parts by Richard Owen. The classification is in part the one used at the present time and was constructed with special attention to details of muscular scars, the hinge, the shell material, and, in some forms, the brachial system. Other important con- tributions during the past one hundred years have been made by J. Barrande (1879) on the Silurian faunas of central Bohemia, Waagen (1879-1895) on forms from the Salt Range in India, Rothpletz in 1886, James Hall, J. M. Clarke, C. E. Beecher, H. S. Williams, E. R. Cumings, P. E. Raymond, C. D. Walcott, F. L. Kitchin, Carl Diener, Charles Schuchert, G. A. Cooper, C. 0. Dunbar, Hertlein and Grant, and numerous others. The majority of the larger monographs and papers contain descriptions of new genera and species from particular areas or formations. Among these are studies on the Guadalupian faunas by G. H. Girty (1908), the Cambrian brachio- pods by Walcott (1912), those of the Kutch Jurassic in India by Kitchin (1900), and the Tertiary forms by Sacco (1902). Other papers deal largely with problems of morphology and phylogeny of special groups or of the class as a whole. Important studies on the development and classification of stages of growth of brachiopod shells were published in 1891 and 1892 by C. E. Beecher, who applied the law of morphogenesis as earlier proposed by Hyatt. The prin- cipal factors used were those of growth and acceleration of development, of me- chanical genesis, and geological sequences of genera and species. Special at- tention was devoted to the study of the embryonic shell or protegulum and its modifications resulting from acceleration, thus showing how the nepionie and neantologic characters are pushed forward and appear earlier in the history of the individual so as to become impressed on the early embryonic shell. As a result of special studies of the pedicle opening Beecher outlined the origin of the deltidium and deltidial plates and developed a classification of stages of growth and decline through the embryonic and larval stages. E. R. Cumings (1903) published a paper on the morphogenesis of the post- embryonic stages of the genus Platystvophia in which he followed the principles used by Hyatt, Beecher, and Jackson. This was a critical study of the nepionie, neanic, ephebic, and gerontic stages of the genus with a general discussion of the history of the genus and the laws governing its evolution. Two important contributions, by Charles Schuchert and by Schuchert and Cooper, have added greatly to the knowledge of the brachiopods. The first paper (Schuchert, 1897) presents tables of species of North American brachiopods arranged by periods and with a classification consisting of 4 orders, 49 families and subfamilies of which 43 became differentiated in the Paleozoic and 30 were extinct before its close. Thirteen continued into the Mesozoic and 6 are represented by living spe- cies. The structural characters are given for each order and the classification is built upon morphologic and phylogenetic principles. The second paper ( Schu- chert and Cooper, 1932) is a detailed study of the suborders Orthoidea and Pentameridae. The authors consider that the division Orthoidea contains the primary stock from which all articulate brachiopods, including the order Telo- 708 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES tremata, have arisen. The classification shows that for the genetic relationships use was made of all parts of the hard anatomy. In the interval between 1910 and 1932 significant advances were made in the classification of certain Upper Paleozoic brachiopods, with emphasis on the in- ternal structures as viewed under the microscope in thin section, 'in 1910 Ivor Thomas published an important memoir on the British Carboniferous Ortho- tetinae and in 1914 a memoir on the British Carboniferous Producti, in which a special terminology was introduced for the investigation of the shells of species which had been collectively grouped under the name Productus. As a result several species were separated from Productus and placed in four newly created genera. In 1910 Stoyanow placed the species typica with a vertical partition in the pedical valve in a new genus Tschernyscheivia. G. S. Girty, also in 1910, described the species elegans with a transverse partition under the new genus Diaphragmus. However, a detailed study of the original material of Productus productus (Martin) by Muir-Wood in 1928 revealed that that species also pos- sessed a transverse partition and, according to the rules of priority, Diaphragmus was placed in synonymy by Dunbar and Condra in 1932. The discovery made by Muir-Wood resulted in the elimination from the genus Productus of a vast number of species with vertical partitions and the erection of twenty-nine new genera and subgenera by Dunbar and Condra. The genus Productus in its re- stricted usage occurs only in the Lower Carboniferous. Other important contri- butions have been made by Stuart AVeller (1914) and Th. Tschernyschew (1902). Pelecypoda and Gastropoda: These organisms have become increasingly abundant in number of genera and species during the course of geologic time, reaching their acme during the Tertiary and Recent. They are thus important to the geologist concerned with the later geologic periods. The advances made in the science of conchology have been intricately associated with the investiga- tions of fossil shells, although the classifications are based largely on the mor- phology of the soft parts. In general, paleontologists have adopted some of the broader groupings of families and subfamilies used by zoologists, but the de- scriptions of genera and species deal almost entirely with the morphological characters of the hard parts. By the middle of the nineteenth century there had been described and illus- trated a vast number of genera and species, both fossil and living, and to enu- merate all the important authors would exceed the limited scope of this review. In the first half of the century the works of Lamarck on the Tertiary mollusks of the Paris Basin and the early, work of Paul Deshayes (1824-1837) presented a classification and laid the foundation for further molluscan research in both Europe and America. Other contemporaneous investigators included the G. B. Sowerbys (father and son), Schlotheim, and Goldfuss. By the middle of the century many investigations were undertaken which resulted in monographic studies of special groups of mollusks as well as of faunas obtained from particu- lar formations. Among the important publications concerned with the morphology and clas- sification of molluscan groups is that of P. Pelseneer (1906) which devotes spe- cial attention to the gills. These characters can be used only with living forms but the groupings of genera based on that kind of information compares well with other classifications founded on a study of the shell characters. In 1884 WEAVEk: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 709 M. Neumayr investigated the hinge structure of bivalve shells and proposed a classification based on the morphological characters of the teeth on the hinge plate. H. Douville in 1912 grouped the Pelecypods on the basis of adaptive radia- tion into a system, proposing three branches depending on the mode of life to which the animal was accustomed. Investigations of both living and fossil mol- lusks by W. H. Dall brought about a classification of the Pelecypoda in which all characters of the shell were considered but with special emphasis on the de- tails of the hinge plate. This grouping of families and genera was used in the English translation of von Zittel's Handhuch der Palaeo7itologie in 1895 and in 1913 (Eastman, 1913), a grouping which, with many modifications, is in general use at the present time. Other notable contributions to the morphology of spe- cial groups of Pelec^'pods and Gastropods are those of R. T. Jackson (1890) on the phylogeny of Pelecypods, F. Bernard (1895-1897) on the morphology of the pelecypod shell, Charles Deperet and F. Roman (1902) on Neogene Pectens, A, W. Grabau (1904) on the phylogeny of Fusus and its allies. Several students of Mollusca have published reference books and monographs which are widely used by paleontologists engaged in the systematic description of fossil pelecypods and gastropods. Among these are H. and A. Adams in 1858, R. A. Phillipi in 1853, Tryon and Pilsbry's Manual of Conchology (1879-1898), J. C. Chenu in 1859, and F. A. Quenstedt in 1881. The many publications by M. Cossmann (1895-1925), including his thirteen-volume work Essais de paleo- conchologie comparee, have been of fundamental importance for classification of gastropod genera. The "Gastropoda" section of the Handhuch der PalaozooJogie (incomplete) by AV. Wenz, 1938-1944, is an up-to-date and extremely valuable treatment of this class of Mollusca. The Handhuch der systematischen Weich- tierkunde, by Thiele, is important for a comparison of the use of soft and hard parts of classification. One of the more recent contributions is Tertiary Faunas, by A. M. Davies (1934-1935). This work considers genera of Foraminifera, Echinoidea, Pelecypoda, Gastropoda, and some vertebrates characteristic of the Tertiary throughout the world and presents the morphological characters that distinguish genera, along with their stratigraphic range and geographical distri- bution. The early half of the twentieth century saw the publication of a very extensive literature by W. H. Dall and P. Bartsch on both fossil and living mol- lusca. These works deal with faunas of particular areas or with biologic groups. A very extensive literature in which fossil mollusks are described deals with faunas of particular formations, with emphasis on stratigraphic problems. The importance of geology to the world-wide growth of the oil industry has stimu- lated research in stratigraphy and in the use of fossils to establish the time sequence of faunas. The result has been the publication of many papers with lists of faunas and the occasional description of new species. Some of the more significant contributions of a purely scientific character during the past one hundred years are those of James Hall on the Paleozoic Mollusca of New York State, S. V. Wood on the Crag Mollusca of Great Britain (1851-1861), Morris and Lycett on the Great Oolite (1850-1863), J. Barrande on Silurian mollusks (1852-1899), Pictet and Campiche (1855-1872) on the Cretaceous Molluscs of Switzerland, and F. A. Quenstedt on the Jura in 1858. Papers dealing with the Mesozoic are those of A. Bittner (1895) on a revision of the Pelecypods of St. Cassian; W. H. Hudleston (1887-1896) on British 710 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Jurassic Gastropods; F. Stoliczka (1868) on the Cretaceous Gastropods of Southern India; H. Wood (1899-1913) on the Cretaceous Pelecypoda of England; L. Waagen (1907) on Lamellibranchs from the Alpine region; A. P. Pavlow (1907) on the Aucellas of Russia; and F. L. Kitchin (1903) on the Jurassic Pelecypoda from the Kuteh area of India, The late Miocene and Pliocene brackish and freshwater faunas of the Balkan Peninsula, South- ern Russia, and the Caucasus have been studied by S. Brusina (1884), N. A. Andrussov from 1897 to 1912, and K. Krejci-Graf and Wenz (1931) who have directed attention to a succession of four nonmarine facies in a series of basins extending from Austria eastward into southern Russia and Asia Minor. In 1905, A. D. Archangelski described the Paleocene faunas in the Saratov area of eastern Russia and their relationship to faunas of similar age in AVestern Europe. A few of the more significant contributions which have aided in making known the rich Tertiary faunas are those of E. Beyrich (1853-1856) on the North German Tertiary; C. L. F. von Sandberger (1858-1863) on Mollusca from the Mainz Basin; Hoernes and Auinger (1879-1891) on the Tertiary faunas of the Vienna Basin; S. V. Wood (1871-1877) on the Eocene Bivalves of England; K. Martin (1879-1880, 1891-1922, etc.) on the Tertiary Molluscs of the Dutch Indies; A. von Koenen on both the Cretaceous and Oligocene of Germany; M. Cossmann on the faunas of many formations in France; R. A. Philippi on the Tertiary of Chili; Nagao, Makiyama, and Hatai on the Tertiary Molluscs of Japan; A. Wrigley on the Eocene of England; and W. S. Slowked- witsch on the Tertiary of northeastern Siberia. The faunas as described and illustrated in this last work closely resemble those of the middle and later Ter- tiary in Oregon and Washington. The scientific contributions to molluscan paleontology of the western hemi- sphere during the past one hundred years have been exceedingly great and only a few of the more important can be listed. Among these are those of T. A. Conrad, who made known the occurrence of Tertiary Mollusca in both the eastern and western parts of North America; James Hall and J. M. Clarke on the Paleozoic mollusks of New York State and the upper Mississippi Valley; F. B. Meek on the Cretaceous mollusks of the Rocky Mountain region in 1876; C. A. White on nonmarine mollusks in 1883; E. 0. Ulrich on the Silurian of eastern North America in 1897; T. W. Stanton on the Cretaceous; and W. H. Dall, H. A. Pilsbry, Julia Gardner, C. W. Cooke, Wendell P. Woodring, W. C. Mansfield, Katherine V. W. Palmer, W. B. Clark, G. D. Harris, and others in many papers on the Tertiary mollusks of the Atlantic and Gulf Coastal Plains and Caribbean regions during the last fifty years. On the Pacific Coast the monumental works of W. M. Gabb and his associates from 1864 to 1869 made known the molluscan faunas of the Jurassic, Creta- ceous, and Tertiary. Near the end of the nineteenth century important papers were contributed by T. W. Stanton and J. C. Merriam on the earliest Tertiary Martinez fauna and its relation to the uppermost Cretaceous. At the turn of the century an important paper on West Coast Cretaceous faunas, including pelecypods and gastropods, was published by F. M. Anderson under the auspices of the California Academy of Sciences. His later papers dealing with the same subject have recently been published by the Geological Society of America. Nu- WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 711 merous papers appeared between 1915 and 1913 by B. L. Clark on the Tertiary mollusks of the Pacific Coast. Contemporaneously, the faunas of the Eocene were described by R. E. Dickerson (1914) and in 1925 the fauna of the type Tejon was described by F. M. Anderson and G. D. Hanna. In 1926 and 1930 two im- portant monographs involving a detailed morphologic and systematic study of Gabb's types from California were published by Ralph Stewart. This work set the standard for more critical reviews of the generic nomenclature of fossil mol- lusks in many of the investigations which followed. A monograph dealing with a detailed systematic study of the mollusks of the Pliocene and Pleistocene of California was published by U. S. Grant IV and Hoyt Rodney Gale in 1931. Investigations of the species of particular genera such as Acila, Nucula and Yoldia, undertaken by TI. G. Schenck at Stanford University during the second quarter of the century, pointed out the desirability of research on special generic groups. A very valuable catalogue was published by A. Myra Keen and Herdis Bentson in 1944 on all of the known Tertiary molluscan species in California. Cephalopoda: The Cephalopods represented by both living and fossil groups were studied, described, and classified in different ways during the first half of the nineteenth century, but the vast literature which has accumulated concern- ing this class during the past one hundred years has revealed a fairly clear knowl- edge of their morphologic relationsliips. The living Nautilus, the cuttlefisli, and Foraminifera were placed in the class Cephalopoda in 1798 by Cuvier and con- sidered distinct from all other mollusks. In 1801 Lamarck noted the differences in the suture lines between Animoiiites and Nautilus and in 1825 De Haan clas- sified the known genera under three families — Ammonitea, Goniatites, and Nau- tilea. Owen in 1832, in a paper on the soft anatomy of the genus Nautilus, pointed out its relation to the Cephalopoda and divided that class into two orders, Tetrabranchiata and Dibranchiata, placing tlie living genus in the former. Von Buch, in papers published between 1829 and 1849, divided the Cephalo- pods into the Nautilidae and Ammonitidae largely on the position of the si- phuncle, and separated the Ammontidae into three sections — Goniatites, Cera- tites, and Ammonites. He introduced technical names for the different parts of the suture lines and used these, along with the varying shape and decoration of the shell, for the establishment of fourteen families. Observations made on the shells of many genera from the Paleozoic through to the end of the Mesozoic showed a progressive complication of the suture lines prophetic of future phylo- genetic investigations. The new avenues of approach for the study of Cephalo- pods formed the groundwork for this type of research between 1850 and 1950 and resulted in the appearance of an extensive literature concerning the phylo- genetic relationships of Paleozoic and Mesozoic genera. Among the more important contributions to the study of the Cephalopoda at the opening of the second half of the last century were the works of F. A. Quenstedt on Der Jura in 1858 and Die Ammoniten des Schivdhischen Jura from 1885-1888. The monographs of Pictet and Campiche from 1858 to 1864 contain descriptions of the Cretaceous Cephalopods of Switzerland and those of J. Barrande (1852-1889) the Silurian Nautiloids of Bohemia. The morpho- logical studies by Suess (1866) showed that, in addition to the details of the suture lines, the variations in the size of the chambers and the shape of the aper- ture were of importance in classification. With the use of these additional char- 712 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES acters he introduced the new generic names PJiylloceras, Lytoceras, and Arcestes. Contemporaneously in North America new methods were advocated by Alpheus Hyatt (1872) for the study of Cephalopods and by 1869 the earlier nomencla- ture of families was abandoned and a new system erected, which was founded largely on phylogenetic considerations. This early work was followed by nu- merous papers elaborating on the principles outlined, with the description of many sharply defined genera. Among the more important of Hyatt's papers was his "Genera of Fossil Cephalopods" in 1884 and the Genesis of the Arietidae in 1889. He introduced the method of study involving a detailed investigation of the successive whorls back to the initial chamber or protoconch and thus at- tempted to unravel the phylogenetic development in terms of the ontogenetic, thereby opening up a new line of research which has been followed by later students. These methods of attack were followed by M. Neumayr on the Am- monites of the Alps and North Germany from 1871 to 1881; Ed. Mojsisovics (1873-1876) on the Triassic of the Alps and on the Upper Triassic of the Hima- layas in 1896; W. Waagen (1879-1895) on the Cephalopods of the Salt Range, India; and K. A. von Zittel on the Stramberger Beds (1868) and on the Titho- nian of the Alpine region in 1870. Neumayr founded his classification of Ce- phalopods on a consideration of direct or close relationship of genera in the line of descent. Others who have added to the information concerning Cephalopods are Thomas Wright (1878-1885) on the Ammonites of the Lias of Great Britain; J. F. Pompeckj (1893-1896) on the Ammonites of Schwabia; A. Karpinsky (1889) on the Permian Ammonites of Russia; W. Branco (1880-1881) on the development and history of fossil Cephalopods; G. G. Gemmellaro (1887-1899, 1904) on the Ammonites of Sicily; and A. Fusini (1897) on the Liassic Ammo- nites of the Appenines. From 1895 to 1919 C. Diener described the Ammonites of the Himalayan region and published papers on the environment, geographic distribution, su- tures, and living chambers of Ammonites. In 1903 R. Hoernes discussed prob- lems of ontogeny and phylogeny. The North German Ammonites were described in 1902 by A. von Koenen and the Jurassic Ammonites of France in 1910 by Dumortier. Papers by S. S. Buckman appeared from 1887 to 1900 on the Am- monites of the Lower Oolite of Great Britain and his monumental work on the Yorkshire Ammonites in 1909. The fine discrimination of species in this last work has been of fundamental importance to Jurassic stratigraphy. Notable con- tributions to the study of Ammonites during the past twenty-five years have been made by L. F. Spath in England and W. Kilian in France. The significant publications of Hyatt in North America were followed by the contributions of numerous authors in the western hemisphere. In 1893 J. M. Clarke discussed the protoconch of OrtJioceras and in 1897 the Lower Silurian Cephalopods of Minnesota. The Ordovician and Silurian Cephalopods were de- scribed by A. F. Foerste in 1921, and other papers dealt with the morphology of Paleozoic genera. These investigations were followed by the important papers of A. K. Miller on nautiloids and Paleozoic ammonites. The families of the Nau- tilidae, Hercoglossidae, and Aturiidae from the Pacific Coast Tertiary were discussed by H. G. Schenck in 1931. In addition to the early papers by Gabb on the Mesozoic Cephalopods of California and Oregon and those of Whiteaves (1876-1903) on the Cretaceous of British Columbia, there appeared in 1902 a WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 713 monograph by F. M. Anderson in which many Upper Jurassic and Cretaceous species from the Coast Eanges were described. Later investigations by this same author on Ammonite faunas and their stratigraphic relationships were published by the Geological Society of America. Investigations on the Carboniferous and Triassic Ammonites of western North America and their relation to similar faunas in other parts of the world were published by J. P. Smith, of Stanford University, in 1903 and 1914. The scientific approach to the solution of his problems was patterned after that introduced by Hyatt. The Cretaceous Ammo- nites of the Upper Missouri region were described by F. B. Meek in 1876. Other important contributions on the Jurassic and Cretaceous Ammonites have been made by J. B. Reeside, Jr. The Jurassic and Cretaceous Cephalopods of southern Texas and Mexico were studied and described by C. Burckhardt from 1906 to 1912 and the Permo- Carbonifierous Ammonites of the Glass Mountains by E. Bose in 1917. Similar paleontologic and stratigraphic studies in northern Mexico have been made by L. B. Kellum and E. W. Imlay during the past fifteen years. The short contri- butions made by d'Orbigny and von Buch during the first half of the past century to the invertebrate paleontology of South America were followed during the next one hundred years by many papers on the Mollusca, including Ammo- nites. The early investigations on the Cretaceous pointed to certain problems to be studied by later authors. Among these investigators were G. Steinmann, 0. Wilckens, A. Ortmann, C. Behrendsen, H. von Ihering, W. Paulcke, T. W. Stan- ton, H. Gerth, P. Groeber, F. Krantz, A. F. Leanza, and E. Feruglio. Arthropoda: The Trilobites, the most primitive group of this phylum, were abundant at the opening of the Cambrian, indicating that their ancestors prob- ably lived in pre-Paleozoic seas, although their remains have not been discovered as yet. The name was introduced by Walch in 1771 (1768-1771, 3:120), and papers by J. W. Dalman in 1827, F. Quenstedt in 1837, Goldfuss in 1843, Bur- meister in 1843, and H. F. Emmrich in 1845 proposed technical names for the different parts of the exoskeleton which were used for classification purposes. These include the number of thoracic segments, the character and position of the facial sutures, the shape and nature of the glabella, the presence or absence of eyes, structural differences, and the ability of the animal to enroll. In 1852 an important monograph by J. Barrande dealing with Silurian Trilobites ap- peared in which all the morphological characters of the exoskeleton were con- sidered and certain phylogenetic relations pointed out. E. Billings in 1870 dis- covered appendages on the ventral side of the genus Asaphus in Silurian rocks and by 1950 similar fossil remains were obtained from Triartlms, Neolenus, Calymene, Ceraurus, and Isotelus. The monograph on British Trilobites by J. W. Salter and H. Woodward, published from 1867 to 1884, is important in that the authors take into consideration the dominant values in ontogeny as a basis for their classification. C. E. Beecher's paper on "Outlines of a Natural Classi- fication of Trilobites," published in 1897, employs the phylogenetic concepts earlier offered by Hyatt and proposes a classification based on the morphogene- sis of all parts of the carapace. He considered the Trilobites especially suitable for the application of the recapitulation theory because of their long history back to the opening of the Cambrian, their generalized structure, and the in- formation available concerning their ontogeny. In an earlier paper on the larval 714 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES stages of Trilobites he described the simple characters of the protapsis and the changes which it underwent during the development of the Trilobite. He showed that in the earlier Cambrian genera this stage is simple but that in the later, more complex genera by a process of acceleration certain characters have been advanced until they appear in the protapsis. He also pointed out that the ven- tral position of the free checks in the earliest larval stages of all except the highest Trilobites is evidence of low rank and for this group he proposed the name Hypoparia. Of the remaining Trilobites, those in which the free cheeks include the genal angles, he placed in the order Opisthoparia and those in which the sutures cut the lateral margins of tlie cephalon he designated Proparia. These three orders, with some additions and refinements, are in general use at the present time. Further evidence was presented to show that the eyes have mi- grated from the ventral side over the margin and then posteriorly across the cephalon to their adult position. Other changes were noted in the character of the glabella and the segments of the pleura. Beecher emphasized the erroneous earlier interpretations of the Trilobites as closely related to the living Limulus. They lack the operculum of the Limulus and possess primitive crustacean af- finities in their protonauplius larval form, slender jointed antennules, the hypo- stoma and metastoma, five pairs of cephalic appendages, and the biramel charac- ter of the limbs. H. M. Bernard in his paper on the systematic position of the Trilobites in 1895 concluded that the crustaceans originated by the bending under to the ventral side of the anterior segments of an ancestral carnivorous annellid. Other significant papers have been published by C. D. "Walcott on Cambrian Trilobites from 1881 to 1916, in which many new species have been described and illustrated. One of these in 1911 is devoted to a discussion of the Cambrian species of China. The Eurypteriids of the Middle Paleozoic were studied by several authors in Europe and North America. F. Koemer in 1848 pointed out their relation- ship to the living Limulus. Other contributions dealing with this group are those of J. M. Clarke and R. Euedemann, who in 1912 described the forms from New York State. The Ostracods, because of their importance in stratigraphic studies, have been investigated by C. I. Alexander (1933), and by R. S. Bassler and B. Kellett (1934), and many others. The fossil Decapods from both the east and west coasts of North America and Central America were described by M. J. Rathbun from 1918 to 1935. Fossil insects occur in certain rocks where conditions for preservation were favorable and were described about one hundred years ago by E. F. Germar from Carboniferous formations near Halle, Germany, and by C. Brongniart from rocks of similar age at Commentry. The Lithographic Shales at Solenhofen also have furnished well preserved fossils, which w^ere described by Meunier, Oppenheim, and Munster. S. H. Scudder in 1879 published an important paper on Paleozoic cockroaches and later (1886) an index to the known fossil insects of the world. In 1900 he described the insects of the Florissant shales of Colo- rado. Other important contributions during the past fifty years include a re- view of American Paleozoic insects by A. Handlirsch in 1906, a monograph by Petrunkevitch (1913) on terrestrial Paleozoic Arachnida of North America, and numerous papers by R. J. Tillyard, including his contribution (1923-1934) on the evolution of the class Insecta. v/eaver: invertebrate paleontology and historical geology 715 Sponsors of Research and Publication During the past one hundred years research and publication of results have been carried on largely by technically trained people associated with national and state surveys, academies of science, organizations with funds available for special problems, and by the geological and paleontological staffs of universities in all parts of the world. The past three decades have witnessed the growing application of earth science to problems connected with the search for oil and gas. As a result, extensive funds have been available for world-wide research and in many instances for the publication of important scientific contributions. The national and state geological surveys were established for the purpose of making known information concerning the natural resources in the rocks and the nature of the problems for their utilization. Many of these organizations have devoted their energies to the investigation and publication of problems con- nected with the direct application of geology and paleontology to the develop- ment of mining in its broadest sense. Others have considered the furtherance of research associated with the more purely scientific phases of paleontology as an important function. A large number of these organizations were founded before 1850. Historical Geology 1850-1950 Pre-Cambrian time: Eocks of pre-Cambrian age have a wide areal distribu- tion involving perhaps twenty per cent of the continents and presumably under- lie as a basement complex all the Paleozoic and later formations. Since 1850 they have been studied extensively in eastern Canada and in the area of the Great Lakes. Over wide areas these rocks have been divided into two broad groups or systems, which are separated by a profound unconformity. The lower division usually consists of granites, gneisses, and highly metamorphosed sedi- mentary rocks and volcanic products : the upper of metamorphosesd and unmeta- morphosed rocks, including slates, quartzites, graywackes, schists, gneisses, and eruptive rocks. Such materials are well exposed and have been studied in detail by many authors in northern Scotland and described as the lower, or Lewisian, and the upper, or Torridonian, systems. Giimbel proposed a similar twofold divi- sion for the basement rocks in Bavaria and Bohemia and, as in North America, referred to the lower series of gneisses and granite as Archaean and the upper gneisses, schists, limestones, and shales as Algonkian. A similar classification was adopted in 1905 by a committee of thirty-five geologists under the direction of Michel Levy for the geological map of France. The pre-Paleozoic rocks of Scandinavia have a wide areal distribution and, after investigation by Torne- bohm, were divided into two series, as in Great Britain. Sederholm in 1907 described an upper, fourfold division of metamorphosed sedimentary rocks as Algonkian; unconformably beneath these were a lower series of granites and gneisses and an upper one of metamorphosed sedimentary rocks. The investiga- tions of von Richthofen in China, followed by those of Bailey Willis and Eliot Blackwelder in 1907, led to a classification somewhat similar to that in the Great Lakes area. In a general way the pre-Cambrian rocks of Western Australia, Africa south of the Sahara, northeastern South America, India, Siberia, and Russia have a similar representation. Certain of the lower formations in Russia 716 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES have an age of 1,852,000 years, as indicated by the study of radioactive distinte- gration products. Investigations of the rocks along the St. Lawrence Kiver Valley in 1843 by Sir William Logan, the first director of the Geological Survey of Canada, re- sulted in the subdivision and differentiation of the pre-Cambrian and the group- ing of the granites and gneisses under the term Laurentian. Later his studies were extended into the area north of Lake Huron where he found a series of slates, quartzites, and conglomerates containing pebbles derived from the under- lying Laurentian granites. He called these rocks Huronian, from their occur- ence on the northeast side of the lake. He recognized a third series of still younger volcanic rocks containing copper and interbeded sedimentary rocks which he considered as a part of the Huronian. Later, in 1876, these rocks were named Keweenawan by Brooks. These studies were followed by detailed inves- tigations of particular areas by Dawson, Bell, Coleman, Collins, and Barlow in Canada and by Van Hise, Leith, Irving, Lawson, Petti John, and many others in the United States. The areas involved included Newfoundland, Nova Scotia, New England, parts of the Appalachian region, Montana, the Grand Canyon area in Arizona, the Llano area of Texas, and many parts of the Cordilleran region. The term Archaean system was defined by Dana in 1872 so as to apply to all pre-Cambrian rocks. Extensive outcrops of greenstones and green schists in the Lake of the Woods area in Canada, which overlie the younger intrusive Lau- rentian granites earlier described by Logan, were studied by A. C. Lawson in 1885 and named the Keewatin series. Lawson also defined the Coutchiching series, consisting of mica schists that were originally sediments, well exposed in the Rainy Lake area and believed that these rocks underlay the Keewatin. He later observed that a thick accumulation of slaty shales, with a basal conglom- erate consisting of boulders derived from the Laurentian granites, rested upon the old intrusive rocks and that these in turn were invaded by a later granite. Lawson named these sedimentary rocks the Seine River series and applied the term "Algoman granite" to the later intrusives. Other rocks of similar age in another region were named the Timiskaming series. Thus there were recognized two periods of batholithic intrusion prior to the deposition of the Huronian sys- tem. Collins, in 1922, determined that a third intrusive interval occurred after the accumulation of the Keweenawan volcanics and was accompanied by strong mountain-making movements which brought pre-Cambrian time to a close. The downwarping of an extensive peneplain carved out of the Algoman Mountains formed an area for the deposition of the Huronian sediments which have been defined as the Bruce, Cobalt, and Animikie formations. North of Lake Huron the lower beds consist of nearly twenty thousand feet of coarse sand- stones and conglomerates containing striated boulders, which were interpreted by A. P. Coleman in 1908 as a tillite indicative of an early ice age. The Ani- mikie series was named in 1873 by T. S. Hunt, of the Canadian Geological Survey, and consists of metamorphosed and unmetamorphosed rocks, including slates, quartzites, graj-vvackes, schists, and eruptive rocks. Originally these were thought to be a part of the Keweenawan series but later this series was found to be unconformable on the Animikie. Because of the great differences of opinion concerning the classification of these rocks a committee of geologists was ap- WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 717 pointed to study them and in 1904 placed the Animikie in the Upper Hiironian and unconforably beneath the Keweenawan. In his earlier work along the north side of the St. Lawrence Valley Logan described a very thick series of metamorphosed limestone and schist, which he named the Grenville series. These sediments contain thick beds of graphite, which are considered to have been derived from some very early organic source. Pre-Cambrian rocks are well exposed in the gorge of the Grand Canyon of the Colorado River and were first made known by Powell in 1875. The L^pper pre-Cambrian sandstones, shales, and limestones were described as being 10,000 feet thick and were named the Grand Canyon series. They are separated by a profound unconformity from the Cambrian above and also from the older horn- blende and micaceous schists and gneisses below, which later, in 1886, were named the Vishnu schist by Walcott. Pre-Cambrian rocks similar to those in the Grand Canyon have been described from the area of the Little Belt IMoun- tains in Montana in the reports of the Hayden Survey in 1872-1873, by Davis in 1886 from the headwaters of Belt Creek, by Peale in 1893-1897, by Weed and Pirsson in 1896, and by "Weed in 1899 from the Fort Benton and Little Belt Mountains quadrangles. These rocks consist of a lower group of greatly meta- morphosed rocks, separated by a marked unconformity from the upper Algon- kian sediments, which were named the Belt series. The latter are -unconformable beneath the Cambrian. The fossil remains of algae, worm burrows, and sponges have been found in the Algonkian rocks but the only evidence of life in the older groups are car- bonaceous slates and graphite. Accordingly, the older rocks have been referred to as Archaeozoic and the younger as Proterozoic, although the United States Geological Survey uses the term Proterozoic for all pre-Cambrian rocks, with the two subdivisions Archaean and Algonkian. Recently, Rankama (1948) has shown that the C^'/C^^ ratios in a number of pre-Cambrian carbon-bearing rocks of Finland are similar to the ratios present in many organic substances and not similar to the ratios in inorganic accumula- tions of carbon. It thus appears that the carbon in these rocks was accumulated by organisms. Rankama concludes that the problematical Corycium enigmati- cum Sederholm from the late Archaean of Finland is a real fossil, probably a primitive alga. This method appears to offer much promise for the determina- tion of the remains of organisms in pre-Cambrian sedimentary rocks. Lower Paleozoic: The controversy concerning the classification of the lower Paleozoic rocks of Great Britain during the first half of the last century con- tinued until 1879 when Lapworth proposed the term Ordovician system for beds previously called Lower Silurian and Upper Cambrian. Thus the lower Paleo- zoic rocks were classed as three independent systems under the names Cambrian, Ordovician, and Silurian. Investigations carried on by the Geological Survey of Great Britain during the past one hundred years show that the Cambrian rocks of the British Isles consist of more than 12,000 feet of sandstones and shale which have been strongly folded, faulted, and in places partially meta- morphosed. The Ordovician formations are best represented in Wales and west- ern England. Important investigations on the lower Paleozoic strata of Bohemia were made by Joachim Barrande and published in twenty-two volumes from 1846 to 1883. 718 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Tlie Middle Cambrian section is one of the most complete in Europe and its abundant fauna is well preserved. The succession of Ordovician beds in the Bo- hemian Basin is representative of the rocks of that system for Europe, and its fossils have been fully described by Barrande. He also made known the richly fossiliferous Silurian limestones of this area, including with them strata which later were considered by Emmanuel Kayser and others to be Devonian. Lower Paleozoic rocks have a wide distribution in the Baltic region and east- ward through northern Russia. The most complete succession of the Silurian formations in Europe occurs on the island of Gotland, where limestones pre- dominate. The Silurian rocks of Sweden and their faunas were studied by An- gelin in 185-4 and, on the basis of Trilobite genera, were divided into seven stages. The Ordovician system in Sweden, as in many parts of the world, is in part composed of shales rich in graptolites, which have been studied by numer- ous paleontologists and correlated with the similar succession in Great Britain. The major subdivisions of the early Paleozoic rocks of eastern North America were studied by Ebenezer Emmons prior to 1850 and in the following years im- portant supplementary contributions were made by James Hall, J. D. Dana, H. D. Rogers, William INIather, C. D. Walcott, and many others. The final re- port of Rogers in 1858 on the geology of Pennsylvania adopted in part the clas- sification earlier proposed in New York State and suggested the idea of the Appalachian trough as a basin of deposition, with a land mass, lying to the east and partly beyond the present coast, as a source of the Paleozoic sediments. The Lower Paleozoic rocks in western Massachusetts and eastern New York, like those in AVales, have been strongly folded, faulted, and partially metamor- phosed; they rest on gneiss, so that the problem of their classification was for many years involved in controversy. Emmons thought these rocks were older than the Upper Cambrian Potsdam sandstone of northern New York and pro- posed for them the name Ta conic system. Numerous investigations during the past one hundred years resulted in an explanation of the Taeonic problem: namely, that moderate uplift during the Ordovician became accentuated near its close, with folding and overthrusting, and a chain of mountains extending from Newfoundland southward to New Jersey was produced. This event has been termed the Taeonic disturbance; in consequence of it the Silurian forma- tions rest unconformably upon the beveled edges of the older rocks. The effect of this disturbance diminished toward the west, where the lower Paleozoic strata are relatively horizontal. The widespread lower Paleozoic formations in the Mississippi Valley and northern Gulf States areas were under investigation by the newly organized state geological surveys during the middle and late nineteenth century. Also, because of the relation of these rocks to the occurrence of oil and gas, particular areas have been studied in great detail during the past fifty years. The name Cordilleran trough has been applied to an area in the Great Basin region extending from Arizona northward into Canada, which during the Paleo- zoic was at times a basin of deposition for great thicknesses of marine Paleozoic rocks. Numerous studies by the U. S. Geological Survey of areas containing mineral deposits have yielded stratigraphic and paleontologic information con- cerning Paleozoic strata and, although different names have been applied to widely separated stratigraphic sequences, a satisfactory correlation of beds is Y/EAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 719 gradually being arrived at. The important monographs on the Cambrian faunas from the Canadian Rockies in Alberta by Walcott have made possible the divi- sion of that system into three series which serve as a standard for comparison with other areas. Surveys made by the geological surveys of Russia and India have shown the presence of extensive areas of early Paleozoic sedimentary marine rocks in north- ern Asia southward into China and also the transgression of the Indian Ocean over parts of western India. Recent investigations show that rocks of early Paleozoic age occur across parts of New Zealand and Australia, in northern Africa, and in the central and western parts of South America. Upper Paleozoic: The Devonian of Great Britain is represented by the Old Red Standstone, which had been made known through the writings of Hugh Mil- ler prior to 1850. When it was realized that these rocks occupied a stratigraphic position between the marine fossiliferous beds of the Cambrian-Silurian and the Carboniferous of Devonshire and Cornwall, they were designated in 1837 as the Devonian system by Sedgwick and Murehison. Since 1850 they have been the subject of many detailed studies by British geologists and now are recognized as of flood-plain and eolian origin. The most complete Devonian sections from both a stratigraphic and faunal standpoint occur in the Rhineland and Eifel areas of western Germany. During the past one hundred years many important contributions have been made to the Devonian rocks and faunas in Russia, cen- tral Asia, and South America. In southwestern Australia Devonian rocks con- sisting of shales and sandstones are reported to have a thickness of nearly 25,000 feet. By 1850 the areal distribution and broad lines of classification of Carboni- ferous rocks were fairly well known. During the past one hundred years many monographs have appeared with descriptions of the faunas and floras and many modifications of stratigraphic classification. In most parts of Europe these rocks have been and still are termed Lower and Upper Carboniferous in contrast to the Mississippian and Pennsylvanian systems of North America. The Carboni- ferous of Germany has been subdivided by Lottner (1868), by H. B. Geinitz (1856), and by F. von Roemer (1870). The Lower Carboniferous beds of West- ern Europe consist largely of fossiliferous limestone, in contrast to carboniferous sandstone and shale in the Upper. Eastward into Asia limestones have been found to prevail. The association of coal, oil, and gas in rocks of Upper Paleozoic age in eastern and central North America has resulted in detailed investigations of these for- mations by State and Federal surveys. The areal and structural geologic maps of large areas of the United States have made known the lithology, thickness, and structure of the Devonian, Carboniferous, and Permian rocks and the classifica- tion of their subdivisions. The Carboniferous system has been recognized by the U. S. Geological Survey, with the Mississippian and Pennsylvanian as sub- systems. These investigations point out the contrast between the strongly folded and faulted beds of the Appalachian area and the slightly tilted strata of the central part of the continent and also with the faultblock structures of the Cordilleran region. A twofold series of uppermost Paleozoic rocks occurs in northern Germany between the Carboniferous and the overlying Triassic beds. The lower part of 720 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES this series consists of red standstones, called the Eothliegendes, and an upper magnesian limestone known as the Zeehstein. Eesting on the red sandstone at the basis of the Zeehstein are black copper-bearing shales, the Kupferschiefer. The Zeehstein also contains deposits of potash salts. Because of the economic im- portance of these materials these rocks were studied in detail in the early part of the last century. Beds of red conglomerate and magnesian limestone in Devon- shire, England, were considered by Conybeare and Phillips to be equivalent to the Eothliegendes and Zeehstein of Germany. Just before the middle of the last century Murchison, de Verneuil, and Key- serling examined the thick series of marls, sandstones, and limestones which rest on Upper Carboniferous beds in the west flanks of the Ural Mountains and pro- posed the name Permian for this system, a term which was immediately adopted in western Europe. In 1874 Karpinsky described beds with a transitional fauna between Upper Carboniferous and Permian and designated them the Artinskian stage. These faunas are known to have a wide distribution from the Arctic Ocean to the Caspian Sea. A complex of Permian and Triassic continental beds occurs in Central and Southern India; these have been studied by W. T. Blan- ford, of the Indian Geological Survey, and named the Gondwana system by Medlicott. These beds have yielded an important fossil flora, including the genus Glossopteris, and many fossil reptiles. The fossils of this series are important because of their widespread occurrence in Australia, Brazil, and South and East Africa and have been used as partial evidence for the proposed continental con- nections called Gondwanaland. The concept of Gondwanaland has been opposed by many geologists and modified by others, notably by Schuchert in his paper on Gondwanaland bridges. Evidence for glaciation during the Upper Carboniferous and Lower Permian occurs in South Africa, India, Australia, Brazil, Uruguay, Bolivia, and the Falkland Islands. The base of the Permian in Central India has been described as consisting of nearly two thousand feet of an old tillite, resting on the striated surface of older beds. These grade upward into sandstones and conglomerates containing Glossopteris. A similar record has been noted by E. H. Schwarz in South Africa, where the Dwyka tillite at the base of the Permian rests on a polished and striated surface, with evidence that the movement of the ice was southward away from the equator. In Australia, T. "W. David reports that the tillite lies on Lower Carboniferous and older rocks and is overlain by coal- bearing sandstones carrying the Glossopteris flora. Largely because of differ- ences of opinion concerning the boundary between the Carboniferous and Per- mian, many European and South African authors have tended to date the late Paleozoic glaciation as Upper Carboniferous. In 1928 Schuchert, after a re- view of the whole Permian problem, concluded that glaciation took place in Middle, and probably late Middle, Permian. He interpreted the climatic change as the result of the Ilercynian orogeny which began in early Carboniferous time and continued periodically through the Upper Carboniferous into Permian time. The Australian geologists T. W. David and C. A. Sussmilch disagreed with Schu- chert's interpretation and in their reply in 1931 gave evidence of six glacial stages, the first two of which were in mid-Carboniferous time, the third in Upper Carboniferous, the fourth in the Lower Permian, and the last two near the top of the Lower Permian. Later studies by A. C. Seward indicate that in Australia WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 721 glaciation began in the Lower Carboniferous, continued into the Upper, and was accentuated in the Permian. In Kashmir the Gangamopteris flora is interbedded with strata equivalent to the Productus limestone. Knowlton considered that this flora originated either in Australia or Antarctica and, with the advance of the ice age, was dispersed northward throughout the southern hemisphere but was prevented from reaching the northern areas by the transverse sea which connected Tethys Basin with the Caribbean Sea and also by the prevailing aridity of northern lands. Prior to 1900 the term Permo-Carboniferous was used widely in North America for sediments now referred to in part as Permian; as early as 1859 it was employed by Meek and Hayden for deposits in Kansas. In 1917 J. A. Udden made known in the Marathon area of western Texas a section of 6,000 feet of dolomites and limestone deposited during the Permian in a sea which occupied large areas of Texas, Kansas, and Oklahoma. This sea was limited on the west by the ancestral Eocky Mountains and on the southeast by the Llanorian uplift. From these lands and from the Arbuckle and Wichita uplifts came the sediments of this age, which are to a considerable extent red beds. Important contribu- tions have been made to the study of these formations by Philip King, of the U. S. Geological Survey. In eastern North America the Paleozoic Appalachian trough was drained and the thick accumulation of sediments folded, faulted, and elevated into the Appalachian IMountains at this time. Triassic: The threefold classification of Triassic rocks of central Germany as the Bunter sandstone, Muschelkalk, and Keuper had been established prior to 1850, largely through the investigations of Alberti. Later each of these divi- sions was subdivided into groups with names based on local variations of lith- ology. The middle marine Muschelkalk member decreases in thickness westward and in England its equivalent, together with the Bunter and Keuper, forms a sequence of continental sandstones, shales, conglomerates, and local beds of gyp- sum and rock salt, which were named the New Red Sandstone. The north Ger- man Triassic became the standard for comparison with the Alpine areas dur- ing the second half of the last century. The threefold division in Germany is not characteristic of the Alpine region, where folded and faulted fossiliferous marine limestones, dolomites, and shales form rugged outcrops extending from Austria westward to the Jura Mountains. Several important monographs on the stratigraphy and faunas of these rocks had been published before the middle of the past century by Lill, H. G. Bronn, Klipstein, Emmrich, Hauer, and von Buch. As a result a partial correlation of the Alpine Triassic with that of northern Germany was made by a comparison of distinctive faunal assemblages. In 1858 F. Hauer, of the Austrian Geological Survey, divided the Triassic succession in the Venetian Alps into seven groups on the basis of the' paleontological sequence. Von Richthof en in 1860 published a work on the Triassic of the South Tyrol, with a full description of the areal dis- tribution, lithology, and tectonic structure of the different formations, and made the suggestion that the limestones of the Southern Alps had been formed by the slow subsidence of reef-building corals. Investigations of the Bavarian Alps by Oppel in 1859 and Giimbel in 1861 led to the recognition of the Dachstein lime- stone and the Kossen formation as the Rhaetic group of the uppermost Triassic. Giimbel, while director of the Bavarian Geological Survey, studied the Alpine 722 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES region in great detail. He proposed the name Vindelic Chain for a former moun- tain range north of the present Alps, extending from north Bavaria westward to the plateau of central France, and accounted in this way for the differences in lithology of the Alpine and Extra-Alpine Triassic. The investigations of E. von Mojsisovics in Austria between 1866 and 1896 emphasized the paleontologic basis for the classification of the Triassic massive limestones in contrast to the earlier divisions founded on lithology. The Triassic rocks of the Himalayas and Salt Ranges of India were studied by von Mojsiso- vics, Diener, and Waagen under the auspices of the Geological Survey of India and have been classified largely on the basis of the divisions established in the Alps. An evaluation of the Permo-Triassic horizons, including that containing the Djulfa fauna in Armenia, has been described by A. Stoyanow. L. F. Spath, in his monograph on the Ammonoidea of the Trias in 1934, has also contributed greatly to the discussion of these problems. The Triassic rocks of eastern North America, extending from Nova Scotia into South Carolina, consist of continental red beds corresponding to the Keuper series of Germany and were named the Newark series. These sediments, ranging from 10,000 to 20,000 feet thick, along with basic lavas, accumulated in down- faulted troughs and were derived from the erosion of the recently uplifted Ap- palachian Mountains. The Triassic continental beds of the Cordilleran basins of western North America were studied for nearly one hundred years hy the United States ex- ploring expeditions and later by the U. S. Geological Survey. They consist of colored shales, sands, and conglomerates, which have been named the Moenkopi, Shinarump, and Chinle formations. Marine members interfinger with the Moen- kopi formation toward the west and in California and Nevada attain thicknesses as great as 20,000 feet. These deposits contain rich ammonite faunas, which were described by J. P. Smith and Alpheus Hyatt in 1905 and by J. P. Smith in 1927. The relation of these faunas to those of the Himalayas and Alps made possible an interpretation of seaway connections which during the Lower Triassic con- nected the Great Basin sea with the Arctic to the north and also with Tethys Basin, as pointed out by Smith. Later the connections were thought to be with the Mediterranean through the Central American portal and a mid-Atlantic archipelago. Again this avenue was closed and a Pacific boreal passage opened. The Upper Triassic faunas of western North America indicate that the connec- tion was once more with the Mediterranean except at the end of the period when boreal faunas again came down from the Arctic. Jurassic: The contributions of William Smith, together with those of Cony- beare and Phillips, made the Jurassic succession in England well known by the middle of the past century. It was classified as the Lias and tlie Ijower, Middle, and Upper Oolite. The early work of Humboldt, Brongniart, Merian, Thur- mann, Dufrenoy, and Elie de Beaumont outlined the general features of the Jurassic rocks of Switzerland, France, and south Germany. Those of south Ger- many were divided into Black, Brown, and White Jura by von Buch, who laid the foundations for the important contributions of F. A. Quenstedt. In these the three main groups were each subdivided into six subgroups and an important section was established in the Schwabian Alps which was extensively used as a standard for correlation. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 723 The monumental work PaJeontologie frangaise, by d'Orbigny (1840-1860) which was partly complete in 1856, presented a classification of the Jurassic rocks and faunas of France, but d'Orbigny still considered that each stage con- tained a specially created fauna, distinct from all others below and above. After a study of the Jurassic in France and England the detailed investigations of Albert Oppel, published in 1858, led to the introduction of the term "zones" for time-stratigraphic units based on the occurrence of certain species which were absent in beds above and below. This work was of fundamental importance for future stratigraphic investigations. The lithologic similarities of the upper- most Jurassic and lowermost Cretaceous rocks in the Alps made it difficult to separate them. Oppel used the term "Tithonian" for the uppermost Jurassic limestones and shales in the northern Alps; and the characteristic Tithonian fauna has made possible the correlation of beds with similar faunas in distant parts of the world. The Upper Jurassic faunas in England and Germany are not everywhere characterized by Alpine species; the upper part of the Tithonian group is now recognized as equivalent to the Portland Purbeck beds, the lower to the Upper Kimmeridgian. The Jurassic rocks of both northern and southern Asia are rich in fossils, which have been compared in important monographs by Waagen, Kitchin, and Noetling, to both the Alpine and northern European faunas. A nearly complete sequence of European faunal zones known in the Jurassic rocks of the Andean trough of South America have been studied by Bodenbender, Steur, Burckhardt, Gerth, Krantz, Jaworski, Leanza, and Feruglio. The absence of marine Jurassic rocks in eastern North America suggests an interval of erosion. During the second half of the past century the Rocky Moun- tain and Pacific Coast areas were investigated by several national exploration surveys, including the U. S. Geological Survey, and by the Geological Survey of California from 1860 to 1869. A shallow western interior sea spread southward from Canada through parts of the Rocky Mountain region into Arizona but was separated from the Pacific Coast embayments by a north and south axial land mass which extended northward through Nevada and eastern California. Dis- cussions of these rocks have been published in the numerous reports of the U. S. Geological Survey, in state reports, and in the geological journals and bulletins. The Jurassic marine formations of the Pacific Coast basins were described in the reports of the Transcontinental Railway Surveys, of the Geological Survey of California, the Geological Survey of Canada, the California Academy of Sci- ences, the present State of California Division of Mines, and by numerous inde- pendent authors connected with the universities. Cretaceous: The many papers, monographs, and maps concerning Cretaceous rocks of western Europe published during the first half of the nineteenth cen- tury made known the principal stratigraphic units and their classification. Among the more important contributors were Leymerie, d'Orbigny, Buvignier, and dArchiac in France; F. von Roemer, Hans Geinitz, and Emmanuel Reuss in Germany and Austria. During the past one hundred years investigations of the Cretaceous of Europe have involved detailed studies of the lithology and faunas of particular areas, the correlation of different stratigraphic units from one area to another and the establishment of faunal zones. Barrois in 1876 attempted to correlate the 724 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Upper Cretaceous succession of England with that of northern France. Coquand carried on detailed investigations in southern France, where the rocks show marked facies variations, and tried to correlate the divisions with those estab- lished by Hebert in the north of France. The regression of the seas late in Juras- sic time left extensive areas of Europe just barely above sea level and upon this surface in southern England was deposited the Wealden formation, with its rich vertebrate fauna. The east-west Vindelican land mass which divided Europe into two east-west basins became flooded as Cretaceous time progressed, producing a succession of overlapping beds from southern to northern France so that by ]\Iiddle Cretaceous time the sea transgressed widely over Europe. Farther east in the Himalaya region of Tethys Basin an extensive series of Cretaceous sediments contained an eastern facies of the Alpine faunas, as described by Victor Uhlig and Stoliczka. Extensive outcrops of folded and faulted Cretaceous rocks, which lie in the great downfold extending w^estward through northern Venezuela and thence southward in the Andean trough from Colombia into southern Argentina and Chili, have been described by the geologists connected with the national surveys of those countries and by independent scientific investigators. The faunas of the successive stages show close relationships to those of Europe. The Creta- ceous faunas of central Argentina have many affinities with those of the Uiten- hage formation of South Africa. Among the more important contributors to these problems are Behrendsen, Burekhardt, Groeber, Windhausen, Gerth, Ort- mann, Krantz, Stanton, Leanza, and Feruglio. The Federal and state survey reports in north America during the second half of the past century present a fairly clear picture of the areal distribution of Cretaceous rocks in the Atlantic and Gulf Coastal Plains, the Rocky ]\Ioun- tain region, and the Pacific Coast. During the past fifty years detailed studies of particular areas have been undertaken by the geologic staffs of oil companies and a part of this information has been published by the American Association of Petroleum Geologists, the Geological Society of America, and other similar organizations. Late in the Jurassic and axial uplift west of the Rocky Mountains was ac- companied by a broad north and south downwarp extending from northern Canada to Mexico and within it were deposited a nearly complete succession of marine Middle and Upper Cretaceous sediments. These formations and their faunas have been described in the numerous reports and geologic folios of the U. S. Geological Survey and other state and ]H-ivate organizations. For many years there was uncertainty concerning the boundary between the uppermost Cretaceous and basal Tertiary, a time of withdrawal of marine seas from the trough and of the sharp tectonic disturbances which accompanied the Laramide Revolution. For some time there were controversies whether the top of the Cre- taceous should be placed at the upper level of the Laramie or Fort Union beds. The land plants in the Laramie at first were thought to be related to the Tertiary but the dinosaurs were distinctly Cretaceous and are not known in the Fort Union beds. Investigations by Earling Dorf in 1940 revealed that the floras of the two formations were distinct, and the plane of demarcation now is placed at the top of the Laramie. The studies made bv the Geological Survey of California from 1860 to 1869 WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 725 showed the existence in the Coast Ranges of thousands of feet of folded marine Cretaceous and late Jurassic sediments, the fossils of which were described by W. M. Gabb and others. Later studies by T. "W. Stanton in 1895 showed that these deposits extend northward to Shasta County. During the past fifty years many important papers have been published by F. M. Anderson, with classifica- tions of the sediments and descriptions of faunas. The Knoxville series, or lower part of the sequence of beds, was assigned by him to the Upper Jurassic and the remainder was considered to represent most of Cretaceous time. Tertiary: By 1850 the broad outlines of classification of the Tertiary of Europe had been established but there were uncertainties concerning the base and top in the Alpine region owing to the complicated structures and the lack of diagnostic fossils. The memoirs of Galeotti in 1837 were followed by those of Dumont in 1849-1852, in which the Belgian Tertiary was subdivided into eleven stages, with the recognition of a series of i)aleontological zones. Lyell placed the lower eight stages in the Eocene, tlie Boldericn in the ^Miocene, and the Diestien and Scaldisien in the Pliocene. Prestwich in 1857, after a study of the Tertiary deposits in the Hampshire and London basins, compared the different formations with those determined in the Paris and Belgian areas and correlated the Thanet sands with the Heer- sien, the London Clay with the Lower Ypresien of Belgium. In 1846 Phillipi, after a study of the Tertiary fossils of Italy, pointed out that a number of living species were present in the Pliocene but this was considered impossible by d'Or- bigny and Agassiz, who regarded the fauna of each stage as an independent unit of special creation. D'Orbigny in 1852 divided the Tertiary deposits of France into four stages, naming them in downward succession as Subapennine, Falunien, Parisien, and Suessonien, the last two being regarded by Lyell as Eocene. This classification has been greatly modified and enlarged during the past one hundred years. The Tertiary deposits and their faunas in the Vienna Basin have been the subject of special study for over a century. After the publications by Bronn in 1837, d'Orbigny in 1846, and Reuss in 1848 these deposits w^ere intensively investigated by Suess, whose important memoir in 1868 presented a detailed de- scription of the Tertiary deposits between the Alps and the Manhart Range. He made known the sequence of beds and their lithologic characters and showed that the Eocene Nummulitic limestone is succeeded in turn by marls, clays, and the Meletta shales, which form an important stratigraphic horizon from the Car- pathians westward through the Alps and into southwestern Germany. Above these in the Vienna Basin are freshwater beds of Eggenburg and Molt, which are succeeded by the brackish-water Cyrena beds and marine clays and lime- stones, which he referred to as the Mediterranean stage. These again are cov- ered by brackish-water sands and clays, which are widely spread in the south of Europe and designated as the Sarmatian stage. These in turn are followed by the Congeria clays and conglomerates of freshwater origin, which he regarded as having been deposited by streams flowing northward from the Alps. He called these last beds the Pontic stage and referred them to the Pliocene. This impor- tant investigation made possible the establishment of a parallel between the Tertiary rocks of the entire Balkan area and the region eastward, in the vicinity of the Black and Caspian seas. 726 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES The Tertiary rocks of Germany occur in the North German Plain, the Rhine- land Basin, and the Schwabian-Bavarian Plateau. The areas of outcrop in north- ern Germany are relatively small and disconnected, and sections with a complete succession of formations are almost nonexistent, rending it difficult to make direct correlations with the well-established stages elsewhere in western Europe. The Tertiary beds in the Maintz Basin were studied by Sandberger (1858-1863) and divided into nine paleontological zones, which were correlated with the stages of Dumont in Belgium. The investigations of Heinrich Beyrich during the middle and second half of the past century shed much light on the North German Tertiary deposits and his detailed studies of the invertebrate faunas led to important refinements in the classification and correlation of the strata. In 1847 he correlated the Sep- tarian clays of north Germany with the Rupelian stage of Belgium because of the identity of fossils. In 1853 in his mongraph on the North German Tertiary Deposits he showed that the fossil species in beds between Magdeburg and Egeln, which he considered Miocene, contained many forms characteristic of older hori- zons and correlated these strata with the Lower Tongrian of Belgium and the Septarian clay with the Rupelian. In 1854 he proposed that these stages, both in Germany and Belgium, should be regarded as an independent series, which he named Oligocene, subdividing this into lower, middle, and upper members. This new unit in the geologic time scale was generally accepted by European geolo- gists and later in the nineteenth and early in the present century was widely used in North America. The Tertiary deposits in the Schwabian-Bavarian Plateau occupy an inter- mediate position between the Swiss and Austrian areas. They were investigated in great detail by Bernhardt Studer, who in 1855 published his Geologie der Schiveiz. He recognized a Jura and sub-Alpine group of deposits, the former consisting of a lower marine division with fossils similar to those in the Mainz Basin and an upper series of freshwater limestones and marls which he con- sidered Upper ]\Iiocene. The sub-Alpine deposits, consisting of freshwater red marls, molasse, sandstone, and beds containing brown coal, extend southwest into the Rhone Valley. The marine fossils obtained from these beds were studied by K. Mayer (1858) who divided the Tertiary deposits into eleven paleonto- logical zones. These stages in ascending order were named Garumnien, Sues- sonien, Londonien, Parisien, Bartonien, Ligurien, Tongrien, Aquitanien, Ilelve- tien, Tortonien, and Astien. The first five were assigned to the Eocene, tlie Helvetien and Tortonien to the Miocene, and the Astien to the Pliocene. In Bavaria the sub-Alpine deposits which lie immediately north of the limestone mountains consist of flysch deposits of Eocene and Lower Oligocene age. These were described in 1861 by Giirabel with a full analysis of the fossils. The peculiar lithologic and paleontologic development of the Tertiary deposits in different geological provinces present great difficulties in making exact correlations from one area to another, and many papers have appeared in the past fifty years which attempt to solve such problems. The fundamental classifications established in Europe have been used as a standard for investigation of the Tertiary in other parts of the globe, but not always with success. The marine Tertiary deposits of North America are confined to the Atlantic and Gulf Coastal Plains and the Coast Ranges of the Pacific slope. The deposits WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 727 on the eastern border are relatively thin and, although slightl.y warped, possess a low seaward dip. In places the thickness is much greater in the Gulf area, where the sediments accumulated in part as delta deposits. On the Pacific Coast thicknesses of as much as 30,000 feet are recorded of both coarse and fine-grained sediments which were deposited in downwarped and faulted basins. Prior to 1850 the general character of the Tertiary in eastern North America was made known through the investigations of T. A. Conrad and others connected with newly organized state geological .surveys. Thick deposits of sandstone, con- glomerate, and shale of fl^uviatile, lacustrine, and alluvial origin which range in age from Paleocene to Pliocene are widely distributed throughout the west- ern interior of the United States and Canada. In the Great Basin area there are several thousand feet of igneous rocks, including tuff, ash, lavas, and intru- sive dikes. There has been much uncertainty concerning the age of the conti- nental deposits but an increasing knowledge of the evolutionary development of the fossil vertebrates, together with evidence from fossil plants, is making pos- sible an acceptable scheme of correlation. The investigations made by W. B. Clark on the marine stratigraphy and paleontology of the Tertiax-y of Maryland and the contributions of E. W. Berry on fossil plants, together with papers by G. D. Harris, W. P. Woodring, Julia Gardner, C. W. Cooke, W. H. Dall, K. V. W. Palmer, and many others have aided in establishing the stratigraphic relationships of the many differing de- posits in the Atlantic and Gulf coastal plains. The importance of these forma- tions in connection with the occurrence of oil and gas has led to the publication of many stratigraphical and faunal papers by geologists on the staffs of the oil companies. The value of foraminifera for determining the age of strata otherwise deficient in diagnostic fossils has made possible a more refined classi- fication of the Tertiary of this area and the correlation of strata which show marked lithologic changes in relatively short distances. The Tertiary deposits in California, Oregon, Washington, and British Co- lumbia were made known during the middle of the past century through the investigations of the Transcontinental Eailway Surveys, the Wilkes Exploring Expedition, and the Geological Survey of California. In California fossiliferous rocks in the vicinity of Tejon Pass were pronounced of Eocene age and ulti- mately became known as the Tejon formation. Farther north in the Coast Ranges other fossiliferous strata thought to be of the same age were also re- ferred to under that name. In 1896 investigations by T. W. Stanton, of the U. S. Geological Survey, and J. C. Merriam, of the University of California, showed that the Martinez beds in central California which at first had been considered Upper Cretaceous should be regarded as Lower Eocene. From this time until 1917 the Eocene of California was classified as Martinez and Tejon. During the early part of the present century the Eocene rocks and faunas were inves- tigated by students and faculty of Stanford University and the University of California and the scientific results were published in the Proceedings of the California Academy of Sciences and the Bulletins of the Department of Geology at the University of California. Many investigations were carried on by the U. S. Geological Survey, with the pul^lication of geological maps and reports. Studies made by E. E. Dickerson, B. L. Clark, Ralph Arnold, C. A. Waring, A. C. Lawson, J. C. Branner, F. M. Anderson, M. A. Hanna, G. D. Hanna, R. 728 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Clianey, and others led to the subdivision of the Eocene into seven stages, desig- nated l)y Clark and Vokes in 1936 as Martinez, Meganos, Capay, Domengine, Transition, Tejon, and Gaviota. These units have been set up largely on evi- dence afforded by molluscan, echinoid, and coral faunas. During the past thirty years parallel classifications have been based on foraminifera and an important standard grouping of faunal zones for the Eocene has been proposed by Boris Laiming. A similar scheme for classifying the Miocene on the basis of fora- miniferal zones was proposed by R. M. Kleinpell and at the present time is widely used on the Pacific Coast. The Tertiary formations of Oregon and AYash- ington correspond with some modification to the classifications set up in Cali- fornia, except that the Oligocene is much better developed. The extensive litera- ture resulting from investigations made by geologists of the oil companies, the universities, the State of California, and the U. S. Geological Survey is laying the foundation for a clear understanding of the geological history of the Ter- tiary period on the Pacific Coast. Much of this information has been brought together and interpreted in the important volume on the geology of California by R. D. Reed (1933). The stratigraphic succession of Tertiary rocks and their faunas in Japan is being made known through many publications in that country. The important paleontological contribution by W. S. Slodkewitsch on the Tertiary of the Kam- chatka Peninsula in northeastern Siberia shows clearly the close relations of these faunas to the Tertiary of Alaska and Oregon and Washington. In the East Indies for over fifty years, beginning about 1880, K. Martin published extensively on the faunas of the Tertiary, determining the ages of the beds largely by means of the percentage of living species present in the faunas. Because of the long distance from the standard European section, correlations with it were difficult. Later in 1931 Leupold and Van der Vlerck elaborated a letter system of classification, from "a" to "h," for the Tertiary of this region, basing it mostly upon the larger foraminifera, and not relating it in detail to the European classification. The continental Tertiary deposits of the AVestern interior region were studied by the geologists of the Federal exploration surveys and later by the U. S. Geo- logical Survey, the American Museum of Natural History, and the geologists and paleontologists connected with universities and state and private museums. The rich collections of fossil vertebrates were at first made known by Cope and Leidy, and later during the present century by Osborne, Matthew, Sinclair, Scott, Lull, Lucas, Loomis, Merriam, Stock, Stirton, and many others. Correla- tion of deposits in widely scattered areas have been based on studies of the evo- lutionary development of fossil mammals. The contributions to the fossil floras of the Tertiary by A. C. Seward, D. H. Scott, F. H. Knowlton, E. ^X. Berry, R. W. Chaney, E. Dorf, and many others have been influential in the correla- tion and classification of the nonmarine Tertiary deposits of North America. Quaternary: The unconsolidated surface deposits between the uppermost Tertiary strata and sediments now in course of deposition in England and the plains of Germany were described by Buckland in 1823 as the Diluvium and were thought to have been carried over the land surface by the waters of the Biblical deluge. In 1839 the name Pleistocene was proposed by Lyell for these deposits. Agassiz (1840), who in his youth had studied the action of living WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 729 glaciers in the Alps, considered that at an earlier time these glaciers had ex- tended out on to the plains, where they formed great ice sheets, and that such conditions had occurred over large areas of the continent. The Diluvium of northern Europe is largely of glacial origin and the Quaternary was thought to have opened as continental glaciation began. In later years Quaternary time was divided into Pleistocene and Recent, the latter representing the interval between the last withdrawal of the ice and the present. Usually this interval has been considered to be about 25,000 years, but carbon^* studies indicate that it may be only slightly more than 10,000 years. In certain parts of the world, as in Greenland and Antarctica, continental ice still persists, whereas in the tropi- cal areas, except at high altitudes, it never existed even during the Pleistocene. Long continued investigations in North America by Alden, Antevs, Cham- berlain, De Geer, Leverett, Bretz, Matthes, Flint, and others have led to the recognition of four glacial epochs, during which the ice sheets advanced south- ward from Labrador and north central Canada halfway down into the United States, and three interglacial epochs, when the ice completely retreated leaving the surface which it had occupied covered with debris carried in and on the ice from northern regions. Each successive glacial deposit rests unconformably upon the much-weathered and eroded surface of the one beneath and it was largely from the interpretation of such data that distinct glacial intervals were recognized. Such studies have made possible an interpretation of the geologic history of the Quaternary. Regions in lower latitudes which were only indirectly affected by glaciation have a history characterized by erosion, deformation, and accumulation of sedi- ments similar to that of the Pliocene and older epochs, as in the Coast Ranges of California where thick deposits of Pleistocene and Recent sands, clays, and gravels of both marine and continental origin accumulated. In many places these strata have been folded and faulted, with evidence of local diastrophism during the Pleistocene. Submarine Investigations : During the past twenty-five years intensive geo- logical studies of the floor of the ocean have been initiated by F. P. Shepard, Maurice Ewing, and Ph. II. Kuenen, and their students. This work has demon- strated that many of the classic concepts of the ocean floor and of geologic processes in the ocean are based on inadequate data and need to be revised. Scholars are now making highly significant discoveries and have demonstrated the existence of great submarine valleys, mountain chains, and escarpments, and have found fossiliferous materials of Cretaceous and Tertiary ages in areas far distant from present land areas. Many important developments affecting historical geology may be expected from this field in the near future. SELECTED BIBLIOGRAPHY AoAssiz, Louis 1833-1844. Recherches sur les poissons fossiles. 6 vols. Neuchatel. 1840. fitudes sur les graciers. 346 pp., 32 pis. Neuchatel. Aluerti, F. a. von 1834. Beitrag zu einer Monographie des Bunten Sandsteins, Muschelkalks und Keupers, und die Verbindung dieser Gebilde zu einer Formation, xx + 366 pp., 2 pis. Stuttgart and Tubingen. 730 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Alexander, C. I. 1933. Shell structure of the Ostracode genus Cytheropteron, and fossil species from the Cretaceous of Texas. Journ. Paleont., 7:181-214, pis. 25-27. Anderson, F. M. 1902. Cretaceous deposits of the Pacific Coast. Proc. Calif. Acad. Sci., ser. 3, 2:1-54. Anderson, F. M., and G. Dallas Hanna 1925. Fauna and stratigraphic relations of the Tejon Eocene at the type locality in Kern County, California. Occ. Pap. Calif. Acad. Sci., No. 9. 249 pp., 16 pis. Andrussov, N. 1897-1912. Die sudriissischen Neogen Ablagerungen. Verh. Russ. K. Mineral. Ges., 34:195-242; 36:101-170; 39:337-495. Angelin, N. p. 1854. Palaeontologia Scandinavica. Fasc. 2, pp. 21-92, pis. 25-41. Leipzig. Antevs, E. 1929. Maps of the Pleistocene glaciations. Bull. Geol. Soc. Amer., 40:631-720. Archangelski, A. D. 1905. Depots paleocenes de Saratov et leur fauna. Mater. Geol. Russia., 22:1-207. Archiac, E. d' 1834-1859. Histoire des progres de la geologie. 8 vols. Arnold, Ralph 1906. Tertiary and Quaternary pectens of California. U. S. Geol. Surv. Prof. Pap., 47:3-146, pis. 1-53. Barrande, Joachim 1852-1899. Systeme silurien du centre de la Boheme. 7 vols. Prague and Paris. Barrell, J. 1912. Criteria for the recognition of ancient delta deposits. Bull. Geol. Soc. Amer., 23:377-446, text figs. Bakrois, C. 1876. Recherches sur le terrain cretace superieur de I'Angleterre et de I'lrlande. Mem. Soc. Geol. du Nord. Lille, l(pt. 1) : 1-232, 1 pi., 2 maps, text illus. Bassler, R. S., and B. Kellett 1934. Bibliographic index of Paleozoic Ostracoda. Geol. Soc. Amer. Spec. Pap., 1:1-500. Beecher, C. E. 1891-1892. Development of the Brachiopoda. Amer. Journ. Sci., ser. 3, 41:343-357; 44:133-155. 1897. Outline of a natural classification of the Trilobites. Amer. Journ. Sci., ser. 4, 3:89-106, 181-207. Bernard, F. 1895-1897. Note sur le developpement et la morphologie de la coquille chez les Lamellibranches. Bull. Soc. Geol. France, ser. 3, 23:104-154; 24:54-82, 412- 449; 25:559-566. Bernard, H. M. 1895. The zoological position of Trilobites. Sci. Progr., 4:33. 1895. Supplementary notes on the systematic position of the Trilobites. Quart. Journ. Geo. Soc, 50:352. WEAVER; INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 731 Berry, E. W. 1920. Paleobotany, a sketch of the origin and evolution of floras. Smithson. Inst. Ann. Rept. (1918), pp. 289-407. Beyrich, E. 1853-1856. Die Conchylien des norddeutschen Tertiargebirges. Zeitschr. Deutsch. Geol. Ges., Vols. 5, 6, 8. Billings, E. 1861. New species of Lower Silurian fossils, Pt. I. Geol. Surv. Canada, p. 5. 1870. Notes on some specimens of Lower Silurian Trilobites, Quart. Journ. Geol. See. London, 26:479-486. BiTTNER, A. 1895. Revision der Lamellibranchiaten von St. Cassian: Arh. K. K. Geol. Reichsanst. Vol. 18. 235 pp. 24 pis. BosE, E. 1917. The Permo-Carboniferous ammonoids of the Glass Mountains, west Texas, and their stratigraphical significance. Univ. Texas Bull., 1762:1-241. Brady, B. H. 1884. Report on the Foraminefera dredged by H.M.S. "Challenger," during the years 1873 to 1876. "Challenger" Rept., Vol. 9. 814 pp., 115 pis. Branco, W. 1880-1881. Beitrage zur Entwicklungsgeschichte der fossilen Cephalopoden Palaeon- tographica, 26(1880) :19-50, pis. 4-13; 27(1880-1881) :12-81, pis. 4-11. Brocchi, Giovanni 1814. Conchiologia fossile subapennine con osservazioni geologiche sugli Appennini, e sul suolo adiacente. 2 vols. Milan. Bronn, H. G. 1834-1838. Lethaea geognostica. 2 vols. Stuttgart. 1848-1849. Index palaeontologicus ... 2 vols. Stuttgart. Brooks, T. B. 1876. Classified list of rocks observed in the Huronian Series, south of Lake Superior. Amer. Journ. Sci., ser. 3, 12:194-204. Brusina, Spiridon 1884. Die Fauna der Congerienschichten von Agram in Kroatien. Beitr. Palaeont. Oesterr.-Ung., 3:125-188. BucH, Leopold von 1834. Ueber Terebrateln, mit einem Versuch sie zu Classificiren und zu beschreiben. Abh. K. Akad. Wiss. Berlin (1833), pp. 43-60. 1837. Ueber den Jura in Deutschland. Abh. K. Akad. Wiss. Berlin (1837), pp. 49-136. 1845. Ueber Cystideen eingeleitet durch die Entwicklung der Eigenthumlichkeiten von Caryocrinus ornatns Say. Abh. K. Akad. Wiss. Berlin (1844), pp. 89-116, 2 pis. BucKLAND, William 1823. Reliquiae Diluviane . . . vii + 303 pp., 27 pis. London. BUCKMANN, S. S. 1909-1930. -Yorkshire Type Ammonites. Vols. 1-7. London. 732 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES BURCKHARDT, CaRLOS 1903. Eeitrage zur Kenntnis der Jura- und Kreideformation der Cordilleren. Palaeon- tographica, 50:1-144, pis. 1-16. Burmeister, H. 1843. Die organisation der Trilobiten. Eng. rev. ed., Roy. Soc. London (1846). vii + 136 pp., 6 pis. Carpenter, H. 1884-1888. Report on the Crinoidea. "Challenger" Rept. Vols. 11 and 26. Chaney, Ralph, and Ethel I. Sanborn 1933. The Goshen flora of West Central Oregon. Carnegie Inst. Wash., Publ. No. 439, pp. 1-103. Clark, B. L., and H. E. Vokes 1936. Summary of marine Eocene sequence of western North America. Bull. Geol. Soc. Am., 47:851-878. Clark, W. B. 1901. The Eocene deposits of Maryland. Pp. 21-92. Maryland Geol. Surv. Clark, W. B., and M. W. Twitchell 1915. Mesozoic and Cenozoic Echinodermata of the United States. U. S. Geol. Surv. Monogr. 54, pp. 1-341, 108 pis. Clarke, J. M. 1893. The Protoconch of Orthoceras. Amer. Geol., 12:112-115. 1897. The Lower Silurian Cephalopoda of Minnesota. Minn. Geol. Surv., Final Rept. 3(2), pp. 761-812. Clauke, J. M., and R. Ruedemann 1912. The Eurypterida of New York. N. Y. State Mas. Mem. 14. 2 vols. Coleman, A. P. 1908. The Lower Huronian ice-age. Journ. Geol., 16:148-158, figs. 1915. The Proterozoic of the Canadian Shield. In Problems of American Geology, pp. 81-161. Yale Univ. Press. Collins, W. H. 1922. An outline of the physiographic history of north-eastern Ontario. Journ. Geol., 30:199-210, figs. Conrad, T. A. 1832-1833. Fossil shells of the Tertiary formations of North America. Pp. 1-56. Philadelphia. 1837. Description of new marine shells from Upper California. Journ. Phila. Acad. Nat. Sci., 7(pt. 2): 227-268. 1842. Observations on the Silurian and Devonian systems of the United States. Journ. Phila. Acad. Nat. Sci., 8 : 228, 280. Conybeake, W. D., and William Phillips 1822. Outline of the Geology of England and Wales, with an Introductory Compen- dium of the General Principles of that Science and Comparative Views of the Structure of Foreign Countries. Ixi + 470 pp., 2 pis., 1 map, text illus. London. COSSMANN, M. 1895-1925. Essais de paleoconchologie comparee. Vols. 1-13. Paris. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 733 CUMMINGS, E. R. 1903. The morphogenesis of Platystrophia ; a study of the evolution of a Paleozoic brachiopod. Amer. Journ. Sci., 15:1-48, 121-136. CUSHMAN, J. A. 1940. Foraminifera. Their Classification and Economic Use. 3d ed. 535 pp., 48 pis. Cambridge, Mass. CuviER, Georges 1811-1812. Recherches sur les Ossemens fossiles. ... 4 vols. Paris. CuviER, Georges, and Alexandre Brongniart 1808. Essai sur la geographie mineralogique des environs de Paris. Pp. 1-278, 1 pi. Paris. Dall, W. H. 1890-1903. Contributions to the Tertiary fauna of Florida. Trans. Wagner Free Inst. Sci. Phila., Vol. 3, pts. 1-6, pp. 1-1634, pis. 1-60. Dalman, J. W. 1827. Om Palaeaderna, eller de sa kallade Trilobiterna. Handl. K. Svensk. Vet.-Acad. (1826), pp. 113-152; 226-294, pis. 1-6. Dana, J. D. 1862. Manual of Geology. 798 pp. Philadelphia. 1872. Green Mountain geology: on the Quartzite. Araer. Journ. Sci., ser. 3, 3:179- 186, 250-256. Darton, N. H. 1928. Red beds and associated formations in New Mexico. U. S. Geol. Surv., Bull. 794, pp. 1-356. Darwin, Charles 1859. The Origin of Species by Means of Natural Selection, ix + 502 pp., 1 pi. London. David, T. W., and C. A. Sussmilch 1931. Upper Paleozoic glaciation of Australia. Bull. Geol. Soc. Amer., 42:481-522. Davidson, Thomas 1851-1886. British fossil Brachipoda. 6 vols. London: Palaeont. Soc. Davies, a. M. 1934-1935. Tertiary Faunas. 2 vols. London: Thomas Murby and Co. Davis, W. M. 1886. Relation of the coal of Montana to the older rocks. 10th U. S. Census, 15:697- 712. Deperet, C, and F. Roman 1902. Monographie des Pectinides neogenes de I'Europe et des regions voisines. Ire partie. Genre Pecten. Mem. Soc. Geol. France, Paleont., 10(26) :l-73, figs., pis. 1-8. Deshayes, G. p. 1824-1837. Description des coquilles fossiles des environs de Paris. 2 vols. Paris. Dickerson, R. E. 1914. Fauna of the Martinez Eocene of California, Univ. Calif. Publ. Bull. Dept. Geol., 8:61-180, pis. 6-18. DiENER, C. 1897. Himalayan fossils, Palaeont. Indica, ser. 15, l(pt. 3) :1 105, pis. 1-13. 734 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES DORF, E. 1940. Relationship between floras of the type Lance and Fort Union formations, Bull. Geol. Soc. Amer., 51:213-235. DOUVILLE, H. 1906. Sur la structure du test dans les fusulines. Compt. Rend. Acad. Sci. Paris, 143:258-261. 1907. Evolution et enchainement des Foraminiferes. Bull. Geol. Soc. France, ser. 4, 6:588-602, 1 pi. 1912. Classification des Lamellibranches. Bull. Geol. Soc. France, ser. 4, 12:419-467. DUJAKDIN, F. 1835. Observations sur les Rhizopodes et les Infusoires. Compt. Rend. Acad. Sci. Paris, pp. 338-340. DUMONT, A. 1849. Rapport sur la carte geologique du Royaume. Bull. Acad. Roy. Belgique, ser. 1, vol. 16, pt. 2, pp. 351-373. DuMORTiER, Eugene 1864-1874. etudes paleontologiques sur les jurassiques du bassin du Rhone. 4 vols. Paris. Dunbar, C. O., and G. E. Condra 1932. Brachiopods of the Pennsylvanian system of Nebraska. Neb. Geol. Surv., ser. 2, Bull. 5, pp. 1-377. Eastman, C. R., ed. 1913. Textbook of Paleontology, Adapted from the German of Karl A. von Zittel. Vol 1. 839 pp., 1594 figs. New York: Macmillan and Co. Emmons, E. 1842. Geology of New York. Pt. 2, Comprising the Survey of the Second Geological District. 437 pp. Albany. Emmrich, H. 1845. Ueber die Trilobiten. Neues Jahrb. Mineral., pp. 18-62, 1 pi. ESTALLON, A. 1859-1861. Etudes paleontologiques sur les terrains jurassiques du Haut-Jura, 2 vols., App. Paris. Feruglio, Egidio 1950. Descripcion geologica de la Patagonia, Republica Argentinica, Direccion Gen- eral de Yacimientos Petroliferos Fiscales. 3 vols., 1107 pp. Fitton, W. H. 1836. Observations on some of the strata between the Chalk and Oxford Oolite in the south-east of England. Trans. Geol. Soc. London, ser. 2, 4:103-400. Fleming, J. 1828. History of British Animals, xxiii + 565 pp. London. Foerste, a. F. 1921. Notes on Arctic Ordovician and Silurian cephalopods. Journ. Denison Univ., Bull. Sci. Labor., 19:247-306, 9 pis. Fromentel, L. E. G. de 1859. Introduction h I'^tude des Sponges fossiles. Pt. 1 (all published) Mem. Soc. Linn. Normandie, Caen, 11 (no. 2) :l-50, 4 pis. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 735 FuciNi, A. 1897. La Fauna del Lias medio del Monte Calvi. Palaeontol. Italica, 2:203-250, pis. 24-25. Gabb, W. M. 1864-1869. Description of the Cretaceous and Tertiary fossils of California, Paleon- tology of California, Geo. Surv. Calif. 2 vols. Galloway, J. J. 1933. Manual of Foraminifera. 483 pp. Bloomington, Ind. : The Principia Press. Gardner, Julia 1935. The Midway group of Texas. Texas Univ. Bull., 3301:1-403, pis. 1-28. Geinitz, H. B. 1842. Charakteristic der Schichten und Petrefacten des sachsischen Kreidegebirges. 116 pp. Dresden. 1856. Geognostische Darstellung der Steinkohlenformation in Sachsen, Die Stein- kohlen des Konigreiches Sachsen . . . Abt. 1. Gemmellaro, G. G. 1868-1876. Studi paleontologici sulla Fauna del Calcario a Terebratula janitor del nord di Sicilia. Nos. 1-3. Palermo. 1887-1890. La Fauna dei Calcari con Fusulina della Valle del Fuime Sosio uella Provincia de Palermo. Giorn. Sci. Nat. e Econ., 19:1-106, 10 pis.; Appendice, 20:9-138, 13 pis. 1904. I Cefalopodi del Trias superiore della regione occidentale della Sicilia. Giorn. Soc. Sci. Nat. Palermo, 24:1-321, 30 pis. Gerth, Hans 1926. Beitrage zur Palaentologie und Stratigraphie des Lias, Doggers, Tithons, und der Unterkreide in den Kordilleren im Siiden der Provinz Mendoza (Argen- tinien):III. Die Fauna des Neokom in der argentinischen Kordillere, geo- logischen Rundschau. 17a: 463-494, pis. 18-19. GiRTY, G. H. 1908. The Guadalupian Fauna. U. S. Geol. Surv., Prof. Pap. 58, 651 pp., 31 pis. 1910. New species of fossils from the Thaynes limestone of Utah. Ann. New York Acad. Sci., 20:239-242. Glaessner, M. F. 1945. Principles of Micropaleontology. 296 pp. Australia: Melbourne Univ. Press. GOLDFUSS, G. A. 1843. Systematische Uebersicht des Trilobiten und Beschreibung einiger neuen Arten. Neues Jahrb. Mineralog., p. 537 et seq., 3 pis. GoLDFuss, G. A., and G. Munster 1826-1840. Petrefacta Germaniae. Vols. 1-2. Grabau, a. W. 1904. Phylogeny of Fusus and its allies. Smithson. Misc. Coll., 44(1417) : 1-192. Grant, Robert 1825-1827. Observations and experiments on the structure and functions of the sponges. Edinburgh Philos. Journ., Vols. 8, 9; New Edinburgh Philos. Journ., Vol. 2. Grant, U. S., IV, and H. R. Gale 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and adjacent regions. Mem. San Diego Soc. Nat. Hist., No. 1, pp. 1-1036, pis. 1-32. 736 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Grant, U. S., IV, and L. G. Hertlein 1938. The West American Cenozoic Echinoidea. Publ. Univ. Calif. Los Angeles Math, and Phys. Sci., 2:1-225, pis. 1-30. GiJEMBEL, C. W. 1858-1861. Geognosticlie Beschreibung des bayerischen Alpengebirges. Gotha, pp. 1-429. 1861. Geognostisclie Beschreibung des bayerischen Aplengebirges und seines Vor- landes. Gotha, pp. 1-950. Hall, James 1843. Geology of New York. Pt. IV, comprising the survey of fourth geological dis- trict, pp. 1-683. Albany. Hall, James, and J. M. Clarke 1898. A memoir of the Paleozoic reticulate sponges constituting the family Dictyo- spongidae. State Geol. Surv., New York, Ann. Kept. (1898), pp. 754-984, pis. 1-47, text figs. Handlirsch, a. 1906. Revision of American Paleozoic insects. Proc. U. S. Nat. Mus., 29:661-820. Harris, G. D. 1896. The Midway Stage. Bull. Amer. Paleont., l(no. 4) :1-125. 1897-1899. The Lignitic Stage. Bull. Amer. Paleont, 2(no. 9) :l-74; 3 (no. 11) : 1-103. Hauer, F. 1858-1859. Beitrage zur Palaeontographie von Oesterreich, bd. 1: (hefte 1-2), pp. 1-64, pis. 1-14. Heinrich, M. 1914. Studien in den Riffkalken des rheinischen Mitteldevons. Zbl. f. Min. usw., pp. 732-736 (also in Journ. Geol., 24:57-60). Hoernes, R. 1903. Zur Ontogenie und Phylogenie der Cephalopoden. Jahrb. K. K. geol. Reichsanst., 53:1-32. Hoernes, R., and M. Auinger 1879-1891. Die Gasteropoden der Meeres-ablagerungen der ersten und zweiten mio- canen Mediterran-stufe in der Oesterreichisch-Ungarischen Monarchic. 382 pp., 50 pis. Vienna. Hudleston, W. H. 1887-1896. A Monograph of the British Jurassic Gastropoda. Palaeont. Soc. London. 514 pp., 44 pis. Hunt, T. Sterry 1873. The geognostical history of the Metals. Trans. Amer. Inst. Min. Eng., 1:331- 345; 2:58-59. Hyatt, Alpheus 1872. Fossil cephalopods of the Museum of Comparative Zoology, Embryology. Bull. Mus. Comp. Zool., 3:59-111. 1884. Genera of fossil cephalopods. Proc. Boston Soc. Nat. Hist., 22:253-338. 1889. A Genesis of the Arietidae. Mem. Harvard Mus. Comp. Zool., 16(3). xi + 238, figs. Jackson, R. T. 1890. Phylogeny of the Pelecypods ; the Aviculidae and their allies. Mem. Boston Soc. Nat. Hist, 4:277-400. 1912. Phylogeny of the Echini. Mem. Boston Soc. Nat Hist, 7:1-491, pis. 1-76. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 737 Jawokski, E. 1915. Beitrage zur Kenntnis des Jura in Sudamerika. Pt. 2. Neues Jahrb. Mineralog. Suppl. 40, pp. 364-456. Karpinsky, a. p. 1874. Geologic explorations in Orenburg district. Abh. K. Mineralog. Ges. St. Peters- burg, ser. 2, 9:212-310. 1889. ttber die Ammoniten der Artinsk-Stufe und einige mit denselben verwandte carbonische Formen. Mem. Acad. Imp. Sci. St. Petersburg, ser. 7, 37:1-104. Keen, A. Myra, and Herdis Bentson 1944. Check list of California marine Mollusca. Geol. Soc. Amer., Spec. Pap. 56, pp. 1-280. Kew, W. S. W. 1920. Cretaceous and Cenozoic Echinoidea of the Pacific Coast of North America. Univ. Calif. Publ. Bull. Dept. Geol., 12:23-236, pis. 3-42. King, W. 1846. Remarks on certain genera belonging to the class Palliobranchiata. Ann. and Mag. Nat. Hist., 18:26-42, 83-94. KiTCHIN, F. L. 1903. The Jurassic fauna of Cutch. Palaeontol. Indica, ser. 9, 3(l):l-87, 14 pis. 1913. The invertebrate fauna and paleontological relations of the Uitenhage series. Ann. So. African Mus., 7:21-250. Kleinpell, R. M. 1938. Miocene Stratigraphy of California. 459 pp., 27 pis. Tulsa: Amer. Assoc. Petrol. Geol. Koenen, a. von 1889-1894. Das norddeutschen Unter-Oligocan und seine Mollusken-Fauna. Abh. Geol. Specialk. Preuss. u. Thuring. Staat., 10 (pts. 1-7) : 1-1458. 1902. Die Ammonitiden der norddeutschen Neocom. Abh. K. Preuss. Geol. Landesanst., Heft 24, pp. 1-451, pis. 1-55. KONINCK, L. G. DE 1842-1851. Description des animaux fossiles qui se trouvent dans le terrain car- bonifere de Belgique. 716 pp., 60 pis. Liege. KOZLOWSKI, R. 1948. Les graptolithes et quelques nouveaux groupes d'animaux du Tremadoc de la Pologne. Palaeontol. Polonica, 3:1-235, pis. 1-42. Krejci-Graf, K., and W. Wenz 1931. Stratigraphie und Palaontologie des Obermiozans and Pilozans der Muntenia (Rumanien). Zeitschr. Deutsch. Geol. Ges., 83:65-163, figs., pis. 1-2. KUNTH, A. 1869-1870. Beitrage zur Kenntniss fossiler Korallen. Zeitschr. Deutsch. Geol. Ges., 21:647-688, pis. 18-19; 22:24-43, pi. 1. Laiming, Boris 1940. Some foraminiferal correlations in the Eocene of San Joaquin Valley, Califor- nia. Proc. 6th Pae. Sci. Congr., pp. 535-568. Lamarck, J. B. P. M. de 1815-1822. Histoire naturelle des animaux sans vertebres. Vols. 1-7. Paris. Lamijeet, J., and P. Thiery 1909-1925. Essai de nomenclature raisonnee des echinides. 600 pp., 15 pis. Chaumont. 738 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Lawson, a. C. 1885. Repoi't on the geology of the Lake of the Woods region, with special reference to the Keewatin (Huronian?) belt of the Archean rocks. Canad. Geol. Surv. Ann. Rept. 1, 151 pp., map. 1913. The Archean geology of Rainy Lake restudied. Canad. Geol. Surv., Mem. 40, pp. 1-115. Leanza, a. F. 1945. Ammonites del Jurassicco superior y del Cretaceo inferior de la Sierra Azul, Mendoza. Anal. Mus. la Plata, n.s., Paleont., Sec. A, 6 (no. 1) :l-99, pis. 1-23. Leymerie, a. 1840. Memoire sur le terrain Cretace du department de I'Aube. Mem. Soc. Geol. France, ser. 1, 4:291-364. Linnaeus, Carl 1758. Systema Naturae. 10th ed. 824 pp. Logan, W. E. 1863. Report on the geology of Canada. Can. Geol. Surv. Rept. Prog, to 1863. 983 pp. Lottner, F. H. 1859. Geognostische Skizze des westfalischen Steinkohlengebirges. Erlauternder Text sur Flotzkarte des westfalischen Steinkohlengebirges, Iserlohn. Loven, Sven 1874. Etudes sur les Echinoides. Handl. K. Sveusk. Vet.-Akad., ll(no. 7) :1-91, pis. 1-53. Lyell, Sir Charles 1830-1833. Principles of Geology. 3 vols. London: Murray. Martin, K. 1879-1880. Die Tertiarschichten auf Java . . . 161 pp., 28 pis., 1 map. Leiden. 1891-1922. Die Fossilien von Java. Samml. Geol. Reichsmus. Leiden. 538 pp., 63 pis. Matthews, W. D. 1915. The Tertiary sedimentary record. In Problems of American Geology, pp. 377- 478. New Haven: Yale Univ. Press. Mayer, K. 1858. Versuch einer neuen Classification der Tertiar Gebilde Europds, Verb. Schweiz. Natur. Wiss. Ges. (1857), 42:164-199. Meek, F. B. 1876. A report on the invertebrate Cretaceous and Tertiary fossils of the Upper Missouri country. U. S. Geol. Surv. Terr., 9:1-629, pis. 1-45. Merriam, Charles 1941. Fossil Turritellas from the Pacific Coast Region of North America. Univ. Calif. Publ., Bull. Dept. Geol. Sci., 26:1-214, pis. 1-41. Miller, A. K. 1938. Devonian Ammonoids of America. Geol. Soc. Amer., Spec. Pap. 14, pp. 1-262, pis. 1-39. Miller, J. S. 1821. A Natural History of the Crinoidea or Lily-shaped Animals, viii + 150 pp., 1 pi. Bristol. Miller, Hugh 1841. The Old Red Sandstone. 275 pp., 10 pis. Edinburgh. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 739 Milne-Edwakds, H., and J. Haime 1850-1854. Monograph of the British Fossil Corals. Palaeont. Soc. 322 pps., 72 pis. 1857-1860. Histoire naturelle des corallaires. Vols. 1-3, Atlas. Paris. MOJSISOVICS, E. VON 1873-1876. Die Cephalopoden der mediterranen Triasprovinz. Abh. K. K. Geol. Reich- sanst., 6:356-835. Morris, J., and J. Lycett 1850-1854. Mollusca from the Great Oolite. Palaeont. Soc, pp. 1-147, pis. 1-15. MOSELEY, H. N. 1877. On the structure of Millepora. . . . Roy. Soc. London, Philos. Trans. 167, Pt. I, pp. 117-135, pis. 2-3. MuiR-WooD, Helen M. 1928. The British Carboniferous Producti Il-Productus (sensu stricto) ; semireticu- latus and longispinus groups. Mem. Geol. Surv. Great Britain, 3:1-217, figs., pis. 1-8. MURCHISON, R. I. 1839. The Silurian system. 2 pts., pp. xxxii + 768 pp., 53 pis., 4 maps. London. MuBCHisoN, R. I., E. DE Vebneuil, and A. Keyserling 1845. The Geology of Russia in Europe and the Ural Mountains. 2 vols. London. Neumayr, M. 1884. Zur Morphologie des Bivalvenschlosses. Sitzber. K. Akad. Wiss. Vienna, 88:385-418. 1887. Die naturlichen Verwandtschaftsverhaltnisse der schalentragenden Foramini- feren. Sitzber. K. Akad. Wiss. Vienna, 95:156-186. Nicholson, M. 1886-1892. A Monograph of the British Stromatoporoids. Palaeont. Soc, 234 pp., 29 pis. London. Ogilvie, Maria 1896. Microscopic and systematic study of madreporian types of corals. Roy. Soc. London, Philos. Trans. (B), 187:83-346. Okulitch, V. J. 1943. North American Pleospongia. Geol. Soc. Amer., Spec. Pap. No. 48, pp. 1-112, pis. 1-18. QppEL, Albert 1858. Die Juraformation Englands, Frankreichs und des siidwestlichen Deutschlands, nach ihren einzelnen Gliedern eingetheilt und vergleichen. 857 pp. Stuttgart. 1859. Die neueren Untersuchungen iiber die Zone der Avicula contorta mit besonderer Beriicksichtigung der Beobachtunger M. Martin's iiber das Auftreten dieser Zone im Dept. Cote d'Or. Wiirttemberg. Naturwiss. Jahresheft., Jahrg-XVI, pp. 315-325. Munich. Gbbigny, Alcide d' 1846. Foraminiferes fossiles du Bassin de Vienne. 312 pp., 21 pis. Paris. 1840-1860. Paleontologie frangaise . . . Vols. 1-6 (until 1891 by various authors). Paris. 1851. Cours ^lementaire de paleontologie et de geologic, Paris. 2 vols, in 3. 382 pp., 227 figs. OSBORN, H. F. 1929. The Titanotheres of Ancient Wyoming, Dakota and Nebraska. U. S. Geol. Surv. Monogr. 55. 2 vols. 740 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Palmer, K. V. W. 1937. The Claibornian Schapliopoda, Gastropoda and Dibranchiate Cephalopoda of the southern United States. Bull. Amer. Paleont., 7:1-730, pis. 1-90. Pavlow, a. p. 1907. Enchainements des Aucelles et Aucellines du Cretace Russe. Nouv. Mem. Soc. Imper. Nat. Moscow, Vol. 17, fasc. 1, pp. 1-93, pis. 1-6. Peale, a. C. 1S93. The Paleozoic section in the vicinity of Three Forks, Montana. U. S. Geol. Surv. Bull. 110, pp. 1-56. 1897. Description of the Three Porks sheet. U. S. Geol. Surv., Geol. Atlas, Three Forks Fol. (no. 24). 5 pp., maps. Pelseneer, Paul 1906. Mollusca. In E. R. Lankester, A Treatise on Zoology. Vol. 5, pp. 1-355. London. Petrunkevitch, A. 1913. A monograph of the terrestrial Paleozoic Arachnida of North America. Trans. Conn. Acad. Arts and Sci., 18:1-137, pis. 1-13. Philippi, R. a. 1836-1844. Enumeratio Molluscorum Siciliae cum viventium tum in tellure tertiaria fossilium quae in itinere suo observavit. Berolini. 2 vols. Berol. and Halis Sax. Pictet, F. J., and G. Campiche 1858-1872. Description des fossiles du terrain Cretace des environs de Sainte-Croix, Mat. Paleont. Suisse. Vols. 1-5. Geneva. POMEL, M. 1872. Paleontologie ou description des animaux fossiles de I'Algerie, Fasc. 5 (Spongiaires), pp. 1-256, 36 pis. Oran. POMPECKJ, J. F. 1893-1896. Beitrage zu einer Revision der Ammoniten des Schwabischen Jura. Ver. Vaterl. Naturk., Vol. 49; Vol. 52 (178 pp., 12 pis.). Powell, J. W. 1875. Exploration of the Colorado River of the "West and Its Tributaries, xi + 291 pp. Washington. Prestwich, Joseph 1857. On the correlation of the Eocene Tertiaries of England, France and Belgium. Quart. Journ. Geol. Soc, 11:206-246. QUENSTEDT, F. A. 1837. Beitrage zur Kenntniss der Trilobiten mit besonderer Riicksicht auf ihre bestimmte Gliederzahl. Wiegmann's Archiv. Naturgesch., ser. 3, 1:337-352. 1858. Der Jura, 842 pp. Tubingen. 1885-1888. Die Ammoniten des Schwabischen Jura. 1140 pp. Stuttgart. Ran KAMA, Kalervo 1948. New evidence of the origin of pre-Cambrian carbon. Bull. Geol. Soc. Amer., 59:389-416. Rauff, H. 1893-1894. Palaeospongiologie. Palaeontographica, 40:1-346, pis. 1-17; 41:221-271, pis. 18-24. Raymond, P. E. 1920. The appendages, anatomy and relationships of Trilobites. Mem. Conn. Acad. Arts and Sci., 7:1-169. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 741 Reed, R. D. 1933. Geology of California. Amer. Assoc. Petrol Geol. xxiv + 355 pp. Tulsa. Reeside, J. B., Jr. 1927. The cephalopods of the Eagle sandstone and related formations of the western interior of the United States. U. S. Geol. Surv., Prof. Pap. 151, pp. 1-87. Reuss, a. E. 1861. Entwurf einer systematischen Zusammenstellung der Foramniferen Sitzb. Akad. Wiss. Vienna, pp. 355-396. Rhumbler, L. 1899. tJber die phylogenetisch abfallende Schalen-Ontogenie der Foraminifereu und deren Erklarung. Verh. Deutsch. Zool. Ges., pp. 162-192. Richthofen, F. von 1860. Geognostische Beschreibung der Umgegend von Predazzo, St. Cassian, und der Seisser Alp in Siid-Tyrol. xii + 327 pp., 4 pis., 1 map. Gotha. Roemer, C. F. 1844. Das Rheinische Uebergangsgebirge. Eine palaontologischgeognostische Dar- stellung. 96 pp., 6 pis. Hannover. ROEMER, F. A. • 1841. Die Versteinerungen des norddeutschen Kreidegebirges. iv + 145 pp., 16 pis. Hannover. 1864. Die Spongitarien des norddeutschen Kreidegebirges. Palaeontographica, 13: 1-64, pis. 1-19. Roemer, Ferdinand 1848. Ueber ein bisher nicht beschreibenes Exemplar von Eurypterus aus devonischen Schichten des Staates New York . . . Palaeontographica, 1:190-193. 1860. Die silurische Fauna des westlichen Tennessee. 100 pp. Breslau. ROEMER, F. VON 1870. Geologie von Oberschlesien. Eine Erlauterung zu der . . . geologischen Karte von Oberschlesien . . . xxiv + 587, xxii [1] pp., 1 pi., text illus. Atlas, 50 pis., 14 maps and sections. Berlin. Rogers, H. D. 1858. The Geology of Pennsylvania, a Government Survey. 2 vols., 1631 pp. Philadel- phia. Rogers, W. B., and H. D. Rogers 1843. On the physical structure of the Appalachian chain . . . Assoc. Amer. Geol. Repts., pp. 70-71 (abst.), 474-531. Rothpletz, a. 1886. Geologisch-palaeontologische Monographie der Vilser Alpen, mit besonderer Brachiopoden-Systematik. Palaeontographica, 33:1-180, 17 pis. Sacco, M. F. 1905. Les Etages et les Faunes du Bass in Tertiaire du Piemont. Bull. Geol. Soc. France, ser. 4, 5:893-916. Salter, J. W., and H. Woodward 1867-1884. A Monograph of British Trilobites, Pt. 1. Palaeont. Soc. London, pp. 1-224, pis. 1-30. Sandberger, C. L. F. von 1858-1863. Die Conchylien des Mainzer Tertiarbeckens. '458 pp., 35 pis. "Wiesbaden. 742 -A CENTURY OF PROGRESS IN THE NATURAL SCIENCES SCHENCK, H. G. 1931. Cephalopods of the genus Aturia from western North America. Univ. Calif. Publ., Bull. Dept. Geol. Sci., 19:435-465, pis. 66-78. 1936. Nuculid bivalves of the genus Acila, Geol. Soc. Amer., Spec. Pap. No. 4, pp. 1-149, pis. 1-18. SCHLOTHEIM, ERKST VON 1820-1823. Die Petrefactenkunde . . . , 3 pts., 52 pis. Gotha. ScHUCHERT, Charles 1897. A synopsis of American fossil Brachiopoda. U. S. Geol. Surv. Bull. 87, pp. 1-464. 1915. Revision of the Paleozoic Stelleroidea with special reference to North American Asteroidea. U. S. Nat. Mus., Bull. 88, pp. 1-311, pis. 1-38. 1928. Middle Permian age of the late Paleozoic glaciation. [Abst.] Bull. Geol. Soc. Amer., 39:208-209. 1932 Gondwanaland bridges. Bull. Geol. Soc. Amer., 43:875-916. ScHucHERT, Charles, and G. A. Cooper 1932. Brachiopoda genera of the suborders Orthoidea and Pentameroidea. Mem. Peabody Mus. Nat. Hist, 4(pt. 1) :l-270, pis. 1-29. Schultze, M. S. 1854. Ueber den Orgauismus der Polythalamien (Foraminiferen). + 69 pp., 7 pis. Leipzig. Scott, W. B. 1913. A history of Land Mammals in the Western Hemisphere. 693 pp. New York: Macmillan and Co. Scudder, S. H. 1879. Paleozoic cockroaches; a complete revision of the species of both worlds. . . . Mem. Boston Soc. Nat. Hist., 3:23-134. 1886. Systematic review of our present knowledge of fossil insects, including myria- pods and arachnids. U. S. Geol. Surv. Bull. 31, pp. 1-128. 1900. Adephagous and clavicorn Coleoptera from the Tertiary deposits at Florissant, Colo. U. S. Geol. Surv., Monogr. 40, pp. 1-148. Sederholm, J. J. 1907. Om Granit och Gneis, deras Uppkomst, Upptradande och Utbredning inom Urberget i Fenno-Scandia. (English summary) Finlands. Geolog. Undersok- ning, Bull. No. 23, pp. 1-110, 8 pis, 2 maps. Sedgwick, A. 1835. [Abst, no title]. Edinburgh, New Philos. Journ., 19:390. 1836. [First use of Cambrian]. British Assoc. Adv. Sci. Rept. 5th meeting, pp. 59-61. Sedgwick, A., and R. I. Murchison 1839. On the classification of the older rocks of Devonshire and Cornwall. [Abst] Proc. Geol. Soc. London, 3(61) : 121-123. Slodkewitsch, W. S. 1938. Tertiary pelecypoda from the Far East, Paleontology of U.S.S.R., Acad. Sci. USSR, vol. 10, pt. 3, fasc. 18, pp. 1-508; fasc. 19, pp. 1-275, 106 pis. Smith, J. P. 1897. The development of Glyphioceras and the phylogeny of the Glyphioceratidae. Proc. Calif. Acad. Sci., ser. 3, 1:105-128. 1903. The Carboniferous Ammonoids of America. U. S. Geol. Surv. Monogr. 42. 211 pp., 8 pis. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 743 Smith, J. P. (Cont.) 1914. The Middle Triassic marine invertebrate faunas of North America. U. S. Geol. Surv., Prof. Pap. No. 83. 254 pp., pis. 1-99. 1927. Upper Triassic marine invertebrate faunas of North America. U. S. Geol. Surv. Prof. Pap. 141. 262 pp., pis. 1-121. Smith, J. P., and A. Hyatt 1905. The Triassic Cephalopod genera of America. U. S. Geol. Surv. Prof. Pap. 40. 394 pp., pis. 1-85. Smith, Joshua Toulmin 1847-1848. Ventriculidae of the Chalk. Ann. Nat. Hist., ser. 1, 20:73-97, 176-191, pis. 7, 8; ser. 2, 1:36-48, 203-220, 279-295, 352-372, pis. 13-16. Smith, William 1815. A Memoir to the Map and Delineation of the Strata of England and Wales, with Part of Scotland, xi + 51 pp., London. SOLLAS, W. J. 1877. On Stauronema, a new genus of fossil hexactinellid sponges. Ann. and Mag. Nat. Hist., ser. 4, 19:1-25, pis. 1-5. SowERBY, James 1812-1845. Mineral Conchology of Great Britain. [Cont. by James de Carle Sowerby.] 7 vols. London. Spath, L. F. 1934. The Ammonoidea of the Trias. Catalogue of the fossil Cephalopoda in the Brit- ish Museum. Pt. 4, pp. 1-521, pis. 1-18. Stanton, T. W. 1895. The fauna of the Knoxville Beds. U. S. Geol. Surv. Bull. 133. 132 pp., 20 pis. 1896-1899. Marine Cretaceous invertebrates. Report of the Princeton University Expedition to Patagonia. Vol. 4, pt. 1, pp. 1-43, pis. 1-10. Stephenson, L. W. 1928. Major marine transgressions and regressions and structural features of the Gulf Coastal Plain. Amer. Journ. Sci., ser. 5, 16:281-298. Stewart, R. B. 1926. Gabb's California type Gastropods. Proc. Phila. Acad. Nat. Sci., 78:287-447, pis. 20-32. 1930. Gabb's California Cretaceous and Tertiary type lamellibranchs. Phila. Acad. Nat. Sci., Spec. Publ. 3, pp. 1-314, pis. 1-17. Stoliczka, Ferdinand 1868-1871. Cretaceous fauna of Southern India: Pelecypoda and Gastropoda. Palaeon- tol. Indica, ser. 1 and 3, Vol. 1; ser. 5, Vol. 2; ser. 6, VoL 3; ser. 8, Vol. 4. Stoyanow, a. 1910. Ueber eine neue Brachiopodenart. Bull. Acad. Sci. of the U.S.S.R. [6], 4(11): 853-855. 1915. On some Permian brachipods of Armenia. Russ. Geolog. Kom., Trudy (Comity g^ologique, Mem.) n.s., No. Ill, pp. 1-95. 1949. Lower Cretaceous stratigraphy in southeastern Arizona. Mem. Geol. Soc. Amer., 38:1-169, pis. 1-26. Studer, Bernhardt 1851-1853. Geologie der Schweiz. 2 vols., illus. Bern. 744 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES SUESS, E. 1866. iJber Ammoniten. Sitzungsber. K. Akad. Wiss., Vienna. Vol. 52, Abt. 1, pp. 71-89. 1868. ttber die Gliederung des Vicentinischen Tertiargebirges. Sitzungber. K. Akad. Wiss., Vienna. 58:265-279. Thomas, Ivor. 1910. Tlie British Carboniferous Orthotetinae. Mem. Geol. Survey Great Britain. Paleontology, 1 (pt. 2) : 83-134. Tbyon, G. W., and H. A. Pilsi:uy 1879-1898. Manual of Conchology. Ser. 1, vols. 1-17. Philadelphia. TSCHERNYSCHEW, TH. 1902. Die obercarbonischen Brachiopoden des Ural und des Timan. Comite Geolo- gique Russ., Memo., Vol. 16, (2). vii + 749 pp., 63 pis. Udden, J. A. 1917. Notes on the geology of Glass Mountains. Bull. Univ. Texas, 1753:3-59. Ulkicii, E. O. 1897. Lower Silurian Lamellibranchiata of Minnesota; Lovs^er Silurian Gastropoda of Minnesota. Geol. Surv. Minn., 3 (pt. 2) : 475-628, 813-1081. Ulkich, E. 0., and R. S. Bassler 1923. Paleozoic Ostracoda, their morphology, classification and occurrence. Maryland Geol. Surv., Silurian, pp. 271-391, 18 figs. Ulrich, B. O., and R. Ruedemann 1931. Are graptolites Bryozoans? Bull. Geol. Soc. Amer., 42:589-604. Van Hise, C. R., and C. K. Leith 1911. The geology of the Lake Superior Region. U. S. Geol. Surv., Monogr. 52, pp. 1-641. Vaugiian, T. W. 1900. The Eocene and Lower Oligocene coral faunas of the United States. U. S. Geol. Surv., Monogr. 39, pp. 1-262, pis. 1-24. VOKES, H. E. 1939. Molluscan faunas of the Domengine and Arroyo Hondo formations of the California Eocene. Ann. New York Acad. Sci., 38:1-246, pis. 1-22. Vredenburg, E. W. 1906. The classification of the Tertiary system in Sind with reference to the zonal distribution of Eocene Echinoidea. Rec. Geol. Surv. India, 34:172-198. Waagen, W. 1879-1895. Salt Range fossils. Palaeontologica Indica., ser. 13, l(l):l-72, pis. 1-6; 2(1): 1-323, pis. 1-40. Wachsmuth, C, and F. Springer 1897. The North American Crinoidea Camerata. Mem. Mus. Comp. Zool., Vols. 20-21. 897 pp., 83 pis. Walch, J. I. 1768-1771. Naturgeschichte der Versteinerungen, zur Erlauterung der Knorr'schen Sammlung. 3 vols. Nuremberg. WEAVER: INVERTEBRATE PALEONTOLOGY AND HISTORICAL GEOLOGY 745 Walcott, C. D. 1898. Fossil Medusae. U. S. Geol. Surv., Monogr. 30. 369 pp., 33 pis. 1912. Cambrian Brachiopoda. U. S. Geol. Surv., Monogr. 51. 872 pp., 104 pis. Weed, W. H., and L. V. Pirsson 1896. Geology of the Castle Mountain mining district [Mont.]. U. S. Geol. Surv. Bull. 139, pp. 1-164, maps. Weed, W. H. 1899. Description of the Fort Benton Quadrangle [Mont.]. U. S. Geol. Surv., Geol. Atlas, Fort Benton. Fol. A (no. 55). 7 pp., maps. 1899. Description of the Little Belt Mountains Quadrangle [Mont.]. U. S. Geol. Surv., Geol. Atlas, Little Belt Mountains. Fol. (no. 56). 9 pp., maps. Weller, Stuart 1914. The Mississippian brachiopoda of the Mississippi Valley Basin. 111. Geol. Surv., Monogr. 1, pp. 1-508, pis. 1-83. WlIITEAVES, J. F. 1876-1903. Mesozoic Fossils. Vol. 1, 416 pp., 51 pis. Geol. Surv. Canada, Ottawa. Williams, H. S. 1895. Geological Biology. 395 pp. New York: Henry Holt and Co. Williamson, W. C. 1858. On the Recent Foraminifera of Great Britain. 107 pp., 7 pis. London. Willis, Bailey 1932. Isthmian Links. Bull. Geol. Soc. Amer., 43:917-952. WiNDHAUSEN, A. '■ 1918. The problem of the Cretaceous-Tertiary boundary in South America and the stratigraphic position of the San Jorge formation in Patagonia. Amer. Journ. Sol., ser. 4, 45:1-53. Wood, H. 1899-1913. A monograph of the Cretaceous Lamellibranchiata of England. 2 vols. Paleont. Soc. London. Wood, S. V. 1848-1882. A Monograph of the Crag Mollusca. 4 vols., suppl. Paleont. Soc. London. 1861-1877. Monograph of the Eocene Bivalves of England. 1 vol. with suppls., 1-182, 1-24 pp., pis. 1-25, A, B. Paleont. Soc. London. Woodring, W. P. 1925. Miocene Molluscs from Bowden, Jamaica. Pt. 1, Pelecypods and Scaphopods. Carnegie Inst. Wash. Publ. 366, pp. 1-222, pis. 1-28. 1928. Miocene mollusca from Bowden, Jamaica. Pt. 2, Gastropods. Ihid. Publ. 385, pp. 3-541, pis. 1-40. Wright, Thomas 1878-1885. Monograph of the Lias Ammonites. 1-503 pp., pis. 1-78. Palaeont. Soc. London. ZiTTEL, K. A. VON 1868. Die Cephalopoden der Stramberger Schichten, . . . Palaeontologische Mitt. Mus. K. Bayer. Staates. Vol. 2, abt. 1, pp. 1-118, atlas. Stuttgart. 1877-1878. Studien iiber fossile Spongien. Abh. K. Bayer. Akad. (Math.-phys. Kl.) Munich, 13:1-154, pis 1-9. PLANT GEOGRAPHY Bij RONALD GOOD University College, Hull, Yorkshire So INTIMATELY is tliG liistoiy of man related to the distrib-ution of plants that it is scarcely an exaggeration to suggest that the study of plant geography must have begun with the first dawnings of man's consciousness of the potentialities of his environment, but today most botanists are content to claim for it a history of more comprehensible length. Some date its beginning from the days of Tour- nefort, who flourished about 1700 and to whom is attributed the first recognition of latitudinal and altitudinal zonation and the way in which these are the reflec- tion of one another. Others call attention to Linnaeus' classification of plant habitats — his Stationes Plantarum — in the Amoenitates. The commonest practice, however, is to regard von Humboldt, the great German naturalist, as the father of plant geography. His travels in South America in the opening years of the nineteenth century resulted in one of the first scientific descriptions of equatorial vegetation, and the recognition of his fundamental contribution (von Humboldt, 1817). Humboldt has the merit of emphasizing that the first chapter of this science was, as it has been in so many others, essentially one of exploration and description. In another sense, also, von Humboldt is a notable landmark. He is the con- necting link between the great voyages of geographical exploration in the latter half of the eighteenth century, among which those of Captain Cook are so prominent, and the series of great scientific expeditions which may be considered to have begun with the voyage of the Beagle from 1831 to 1836. In view of the great distinction of Charles Darwin's subsequent studies in botany one cannot but regret, on reading his account of this voyage, that he was not then more con- cerned with plants. His preoccupation at that time with geology and zoology, an emphasis which later had such profound consequences, is evident, and it was not until a few years later, when the young Joseph Hooker set out in the Erehus on a voyage lasting from 1839 to 1843, that a real botanical milestone was reached. It is not easy now to realize that these two series of expeditions — the great voyages of geographical adventure on the one hand and the great scientific explorations initiated by the voyage of the Beagle on the other — ^were in fact separated by little more than half a century for they seem to belong to different ages. True, these years had been memorable ones and had witnessed the vast liberating forces of which the American War of Independence and the French Revolution were expressions, yet even this does not adequately account for the difference of outlook that distinguished the second quarter of the nineteenth century from the third quarter of the eighteenth. It seems clear enough that there must have been in the latter period a tremendous intellectual leaven at work which was destined in the space of a comparatively few years to lighten the whole body [747] 748 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES of biological tlioiight. This leaven may be described as tlie growing consciousness, not yet expressed in words but evidently present in the minds of many people, that the uninspiring doctrines of biological immutability must soon give way to something more in tune with the spirit of the times and, it may be added, more in accord with a rapidly growing body of observed facts. The spark that ulti- mately fired this tamped charge was, of course, the appearance of Darwin's The Origin of Species in 1859, but it is easy to see now not only that the fuse had been smouldering for years, but that the study of plant geography had made no small contribution to this result. Indeed, it may be said with truth tliat, from the point of view of this sub- ject, the foundation of the California Academy of Sciences in 1853 could scarcely have been at a more auspicious time, for it was followed within the short space of seven years by a series of pul^lications which became classic and which, together, raised the subject of plant geography to a position of special importance and sig- nificance. First came Hooker's essay on the New Zealand fiora (1853), to be fol- lowed two years later by A. L. P. P. de Candolle's Geographie Botaniq^ie Raison- nee (1855), which still remains one of the most considerable of all such works. Then, almost together, came Asa Gray's study of the flora of Japan (1859), the Origin itself (1859), and Hooker's second essay, on the flora of Tasmania (1859). Looking back now in the light of so much after-knowledge it is difficult to recapture the intellectual atmosphere of the earlier eighteen-fifties, when the scientific world was so much smaller than it is now. Such recapture is particularly difficult when based on much of the contemporary literature. De Candolle's book is an instance of this. Here is a closely packed study of his subject of more than a thousand pages, of which the headings might serve almost equally well for a survey of similar scope today and in which the author comments with judgment on almost every aspect of the subject, and yet it is written entirely in what can today only be called the restricted idiom of pre-evolution. Even mutability is admitted, and there are discussed, more than once, the changes which species may come to suffer with the passage of time. But there the curtain falls, and one may search in vain for any recognition of the possibility that what may in due course befall species may itself be the origin of others. It is a remarkable example of scientific thought restrained by dogma. De Candolle's book is in many ways a striking example of one written years before its time. It not only discusses many subjects the potentialities of which are really only now being tested, such, as the order of families in floras or the proportions between monocotyledons and dicotyledons, but, as has been said, it discusses the circumstances surrounding the hypothetical creation of species with considerable acumen. So much emphasis is placed on this aspect of his sub- ject that one almost inevitably wonders whether the author's major premise, which is the supernatural creation of species, can have been more than a piece of traditionalism designed perhaps to ensure that the rest of his work would not be dismissed too summarily. But this does not appear to have been the case. Thistleton-Dyer (1893), in his obituary of de Candolle, suggests that it was partly the influence of ideas about the climatic factors of distribution and partly a somewhat unimaginative quality of mind that made him miss the essential point by so narrow a margin. How near he had been to it he himself fully realized later, and it is pleasant to notice in after correspondence Darwin's great GOOD: PLANT GEOGRAPHY 749 opinion of the Geographie Botanique Raisonnee and his appreciation of its author's generous attitude toward his own theories. Asa Gray's relation to the story of the Origin is different. He was one of the few confidants whom Darwin had kept informed of the gradual development of his own theoretical opinions about evolution. Indeed, one of his frequent letters to Gray, which happened to express some of his ideas particularly concisely and conveniently (as well as to date them), was included as part of the joint com- munication of Darwin and Wallace to the Linnean Society of London by which the theory of natural selection was launched on July 1, 1858. The exact rela- tion of Gray's classic paper on the North American and Japanese floras to the Origin is not simple to gauge now. This relationship was the subject of corre- spondence between the two authors during the preparation of Darwin's great work for the press, and it seems fair to regard Gray's paper as having been written in a form intended to provide possible additional evidence for Darwin's views. On the other hand Gray's formal attitude toward these views of Darwin's was one of studied caution. Although Darwin considered Gray one of his most valuable supporters, this support was tempered by a certain criticism because, it would seem. Gray felt that a judicial attitude of this kind would be more effec- tive aid in the long run than any more spectacular and enthusiastic championing. Hooker's two great papers were rather more profound phytogeographical studies but are perhaps to be thought of as less intimately involved with the trend toward evolution. They were obviously of great importance to it, since the facts that they set forth spoke for themselvess in no uncertain voice. Of course Hooker was even more closely associated with Darwin than was Gray but, to use an apposite simile, the work of the two seems to illustrate convergence rather than strict homology. It is perhaps an interesting commentary on this point that Hooker, at any rate from 1850 to 1856, repeatedly expressed pessimistic opinions about the progress and future of botany as a science, which suggests that he was not altogether fully conscious of how rapidly the renaissance was approaching. Let us hope that some of the pessimisms of today are equally ill-founded. However, we must not allow ourselves to wander farther down the by-ways of evolutionary history, fascinating though these are. What has been outlined has been intended to illustrate how opportune the founding of the Academy was and to call attention to the three great botanists, de Candolle, Gray, and Hooker, who dominated the phytogeographical scene at that time. There is never- theless one other point about this birthday which must not be overlooked here, namely, that it occurred only three years after California had become a state of the Union. This surely is but further evidence, if such were needed, of that intellectual leaven to which reference has already been made. With the coming of evolution the whole meaning of plant geography altered. It has often been said that the real evidence of the truth of the theory of evolution lies, not in this or that array of facts, but in its power as an organizing concept. Without it, the facts appear chaotic; with it, they fall into order to a remarkable degree. Nowhere is this more true than in plant geography, which is so funda- mentally a mass of descriptive fact, and this is one of the reasons why it became to tlie evolutionists one of the most promising and popular aspects of botany. The other and even more important reason was the recognition, under the new conception, of the inevitable relation between time, space, and change. All this 750 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES meant that the world vegetation and its distribution, which previously had been, as it has been expressed, matters for wonder but not for speculation, could now be, and indeed had to be, restudied from the point of view of the new theory, and a vast new field thus opened. Plant geography had, in short, to be trans- lated into the new idiom. In this way the whole subject developed so quickly that only the most prominent features of its history can be noticed in one short paper. It is very obvious from a study of botanical literature in general that the characteristics of different nations and peoples express themselves as much in their methods of scientific inquiry and in their predilection for certain aspects of their subjects as they do in many other ways. A philosopher could probably explain convincingly why it was that the German school of botanists almost at once made the newly opened fields so particularly their own, quickly reaching in them a preeminence which they maintained for many years. If an explanation is to be hazarded here, it is that this next phase of the subject was necessarily one of bringing some sort of order out of an enormous and rapidly accumulating mass of data, and that this was work which called for just those qualities of application, industry, and organization that are such strong German national features. The first, and in some ways the greatest, of these German publications was Grisebaeh's Vegetation der Erde (1884), which first appeared in 1871, then in an enlarged edition in 1884. This may almost be described as the first full-scale attempt to give a coherent single description of the vegetation of the whole world and to classify it floristically, and the best compliment that can be paid it is to say that it is still an extremely valuable source of basic information. Indeed, the most striking thing about it now is how little it has been rendered obsolete by subsequent increase in our knowledge, and one can only be surprised that so authoritative and complete an account could be prepared at that com- paratively early date. Actually, as the preface to the first edition says, the book is a synthesis of studies extending over thirty-five years, and thus is partly pre- Darwinian, as is evidenced by the stress laid on temperature as a distributional factor. This emphasis derives from de Candolle, and is partly Darwinian, as is shown by its clear expression of the evolutionary conception of adaptation to environment. It must also be borne in mind that this book dates from a time when the distinction between floristic and vegetational studies had scarcely begun to be made, and it would perhaps be fairer to regard it as an early essay in what later became distinguished as plant ecology (it contains, for instance, one of the first classifications of growth form), though it also includes much direct information about the spatial distribution of plants. A contemporary study more definitely developmental in outlook was the briefer early work of Engler which is often referred to as the Versuch (1872-1882). The mention of Engler, who was, within the scope of his interests, one of the greatest of all German botanists, brings to mind another gradual divergence of subjects such as is inevitable with the passage of time and the growth of material. The study of plant geography must always rest largely on the devoted work of the taxonomists. In earlier days the two fields were almost parts of one whole, but later taxonomy came to absorb nearly all the energies of its chief practitioners. This is true of both Hooker and Engler, whose careers have many GOOD; PLANT GEOGRAPHY 751 interesting parallels. Bach was early attracted by geographical problems and retained his interest in these throughout his long life. But both later devoted themselves especially to systematics, Hooker's work in this field culminating (in collaboration with Bentham) in the Genera Plantarum (1862-1883) and in a number of floras, of which that of British India is preeminent (1875-1897), Engler's work in his well-known Syllabus (11th ed., 1936), and in the editing of such great undertakings as the Naturlichen Pflanzenfamilien (Engler and Prantl, 1889-1924) and Das PflanzenreicJi (Engler, 1900). Much the same is true, too, of de Candolle, whose energies were later deeply absorbed in the con- tinuance of his father's Prodromus (A. P. and A. L. P. P. de Candolle, 1824-1873) . Drude was perhaps in the more direct geographical succession, and in par- ticular will always be remembered for his Atlas der Pflanzenverhreitung. This was published in 1887 as part of a larger physical atlas and consisted of a short series of excellently produced maps showing the ranges of various important plant elements, both vegetational and floristic, accompanied by a concise explana- tory letterpress. The work of Drude, however, is more generally known from his Handhuch der PflanzengeograpJiie, which appeared in 1890 and contained among other things an improved floristic classification. This, however, though an im- portant book, said little that was entirely new and gives the impression rather of belonging to the end of an epoch. It is very noticeable, in the gradual development of a science, how often progress takes the form of successive pulsations, each giving great impetus to the study for a time but then tending to lose momentum, being replaced in due course by some new intensification along some rather different line. Thus it would seem that by the eighteen-nineties the forward urge provided by Darwinism had begun to work itself out and that some new impulse was due. This came in the form of a concentration upon the relation between the plant and its immediate environment or habitat, a new approach or point of view to which was given the name "plant ecology," or "oecology," as it was first spelled. The first principles of this new discipline, which, as we shall see, has since become the sister of the older plant geography in the stricter sense, were set forth in two books which were, effectively, more or less contemporaneous. These were Warm- ing's Plantesamfund, published in Denmark in 1895 and later translated into the more familiar Oecology of Plants, and Schimper's Pflanzen-Geographie auf Physiol ogische Grundlage (1898), which also was translated into English some years later. There is no doubt that a powerful influence in the hiving off of plant ecology was the reaction against the aridity which had affected much of botany through an overemphasis on formal morphology. It may be said to have been based on two fundamental propositions : that the plant itself is a living organism in close and intimate relation, both functionally and structurally, with the conditions of its environment, and that vegetation is a dynamic complex expressing the same laws of universal change with time as everything else in nature. Schimper himself expressed this idea (loc. eit.) when he said that the problems of plant geography will not be exhausted when the world flora is completely known (a contingency which, strangely enough, he seems to have thought imminent) but will become of a rather different sort and particularly concerned with the explanations of the differences between floras in different parts of the world. 752 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES For vegetation is always developing; floras occupy only a moment of vegetational history; and it is the relation of structure, function, and environment which must be studied. It was very early recognized that plant geography is a matter which involves, in a peculiarly direct way, the fundamental conceptions of both space and time, and that the subject might, therefore, be approached from one or other of these directions or from some combination of both. Thus there has always been in plant geography an underlying triplicity; the swing of emphasis within this provides the background to the history of the subject. Just how the three streams or branches of plant geography should be defined and named has been a matter of considerable argument. Those who are interested in terminolo- gies will find excellent accounts of this by Riibel (1927)' and by Wulff (1943), but it is more convenient here to describe this triplicity in rather freer terms. First, there is the stream in which the main emphasis is the correlation of space and form; this has been the special concern of those plant geographers, like de Candolle, Hooker, Asa Gray, and Engler, who have also been preeminent systcmatists. It may be called the taxonomic stream. Second, there is the stream in which the strongest emphasis is placed upon the historical and developmental aspects of the subject, and this is perhaps most often now called the historical stream. Third, there is the stream in which the two conceptions of space and time are balanced more evenly than in either of the others, namely ecology, which is mainly concerned with the distributional changes, usually by their nature relatively small, resulting from the gradual changes in a mutable environment. The main *point in relation to this analysis is very clear : the coming of Dar- winism shifted the emphasis away from the first stream, where it had in fact been almost wholly concentrated, and distributed it more evenly among all three. For a good many years the full effect of this was not felt because this was a period of reorientation, but once this adjustment had been made, the rapid development of the streams which had been so long held back by the pre-evolu- tionary conception of the cosmas was inevitable. For reasons which we need not attempt to specify too closelj^ but which are certainly connected with the full flowering of the idea of adaptation to environment the ecological stream was the first to break through. So simple an analysis is likely to be too clear-cut to depict the whole truth, and this is certainly so here, especially with regard to the first two streams, between which there has always been a close connection. We may indeed recog- nize two streams, but there is, as it were, a constant interchange of water between them. The ecological stream, however, has much more noticeably scoured its own channel and, although this stream flows alongside the others, there is little actual communication between them. The reasons for this would make a most interesting study, for they are probably not all purely botanical, but this is no place to attempt it. We must content ourselves with the statement that what is now called plant ecology became in a comparatively short time largely divorced from the other aspects of plant geography. There is therefore both reason and excuse for referring only briefly to it here, apart from the fact that, since ecology was unknoAvn as a separate study in 1853, it may formally be considered outside the terms of present reference. It is difficult to mark the exact point at which plant ecology became estab- GOOD: PLANT GEOGRAPHY 753 lished as a separate subject, but it is not unreasonable to think of Schimper's Pflanzen-Geog raphie as belonging to the older dispensation and of Warming's Plant esamf unci as belonging to the new, though in fact this is the reverse of their actual dates of appearance. Apart from these two it was perhaps the work of Flahault, of Montpellier, rather than that of any other man that gave the initial impulse to ecology, which at first consisted largely of the mapping of vegetation, such as he had been doing in France for some years. At all events it was one of his pupils, Robert Smith, who introduced his methods into Britain. But shortly after this Smith unfortunately died and, although his work was carried on by his brother and others, it later met with practical difficulties. Attention then passed, largely under the leadership of Tansley, happily still with us, rather to the analysis of vegetation and the study of the different kinds of plant communities, work in which a very definite stage was reached by the publication in 1911 of Tansley 's Types: of British Vegetation. As might be assumed, a similar and indeed even greater development had been taking place synchronously on the continent of Europe, in which indeed so many names claim recognition that it is almost invidious to make a selection. Among the significant works the following stand out clearly: Schroter's publica- tion (with Frith) of Die Moore der Schiveiz (1904), which was a landmark; Raunkiar's classic study of growth form (1907) ; and Riibel's later work, Pflanzen- geselhchaften der Erde (1930), which is a remarkable study of European plant communities. According to Tansley (1911) Schroter also deserves mention as the first to distinguish between ''synecology," or the study of plant communities in relation to their habitats, and "autecology," or the study of the ecology of single species. In America the growth of the new subject went hand in hand with that in the Old World, as instanced by Hitchcock's OecoJogical Plant Geography of Kansas (1898) and by the Phytogeography of Nebraska by Pounds and Clements (1900), but with a rather greater emphasis on the developmental aspects. By 1899 Cowles had begun to publish on the subject of plant succession, the great later expansion of which under the leadership of Clements (1916) is one of the notable features of American plant ecology. It is interesting to note this differ- ence of emphasis for it is surely indicative of the great distinction between European and American ecological development. European botanists had, of necessity, to work upon a vegetation which could be considered natural only by a considerable exercise of imagination, whereas the American school had as its subject vast areas of country over which the influence of man had scarcely been felt at all. It is not surprising in this circumstance that American ecology developed very rapidly and, in many directions, soon attained a leading position. Although it has been convenient to regard plant ecology as stemming from Schimper and Warming, it needs to be stressed that this really marks the for- mal separation of the subject — its coming of age — rather than its birth, for these were certainly not the first publications written from the ecological stand- point. There were, for instance, the studies of Graebner (1895) and others on the North German heaths in the earlier 'nineties; and, especially, Drude's account in 1890 of the plant formations of Central Europe, which actually incorporated some forms of later ecological nomenclature. Earlier still, in the 'eighties, there were Krasnov's account of some of the Russian steppes (1886), Sargent's 754 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES report on the North American forests (1884), and Christ's book on the Swiss vegetation (1879). Indeed, from almost the earliest days there had been a slowly increasing concern with ecological problems in plant geography, until shortly before the first World War the literature of ecology had grown so great that it became desirable to establish special periodicals to accommodate it, a stage it which we may perhaps consider the subject to have passed, for the time being at least, beyond the purview of this paper. The end of the nineteenth century marked also the centenary of the depar- ture of von Humboldt and Bonpland on their travels in tropical America. Readers will find an interesting account of the development of plant geography up to 1900 in Engler's contribution to the Centenary volume of the Berlin Geographical Society (1899), in which he first traces the beginnings of the subject from the earliest times and then its gradual growth on the fioristic side, region by region. The double deflection, or apparent deflection, of interest which followed the turn of the century, on the one hand toward more narrowly taxonomic work and on the other toward ecology, for a time left the middle stream of plant geography a somewhat feeble one. In this direction, at any rate, the fifteen years or so preceding the war were not among the most remarkable. This diversionary tendency was intensified also by one of the most considerable advances of that time, the growth of the subject of genetics, for, as will be seen, it was not until considerably later that the underlying unity between genetics and plant geog- raphy became perfectly realized. Nevertheless these years were far from being entirely barren. In particular the German school continued to demonstrate its leadership in its chosen fields by the continuation or launching of such great projects as Engler and Drude's Die Vegetation der Erde (1896 — ) — among the volumes of which Harshberger's Phytographical Survey of North America is conspicuous — and Karsten and Schenk's V egetationshilder (1903 — ) which presents so much of interest and importance in the international language of illustration; by such books as those of Solms-Laubach (1905) and Schroter (1912); and by innumerable shorter publications, notably in Engler's Botanische Jahrhiicher. Most of these are on the border line between historical and taxonomic plant geography, but an impor- tant direct contribution to the former was the comparison by Engler of the floras of tropical Africa and of tropical America, a subject which was later to become much more topical. In addition, these years saw the earlier writings of several whose major con- tributions to plant geography were to come after the war, among them Fernald, Merrill, Skottsberg, and Willis, but one of the most important series of writings came from H. B. Guppy. Guppy was not a biologist by professional training ex- cept in so far as he began his career as a naval surgeon, but he was that much rarer thing, a born naturalist and observer, and he made good use of the fortune which took him for many years to what are some of the most interesting parts of the world from the point of view of plant geography. His larger works, namely. Observations of a Naturalist in the Pacific (1903-1906), Studies in Seeds and Fruits (1912), and Plants, Seeds and Currents in the West Indies and the Azores (1917), are perhaps a little voluminous and prolix for ordinary reading but they are unquestionably the work of a mind possessed of unusual descriptive and analytical powers. The second volume of the first-mentioned work, which GOOD: PLANT GEOGRAPHY 755 deals with plant dispersal, will long remain a classic source of fact and commen- tary on that subject and its many related problems. Eather later on, and partly in association with Willis, Guppy turned his attention toward historical plant geography, and two of his papers in this field. The Island and the Continent (1919), and Plant Distribution from the Standpoint of an Idealist (1917-1920), are notable for their penetration and freshness of thought. Again, although the prewar years may not have been very eventful scientifi- cally, it was in this period that the foundations were laid for much of the later progress, especially in the increase of knowledge during the years in two fields relating to the distant past. First, it was a time of great activity in paleobotany, and although the more spectacular expressions of this centered in epochs too re- mote to interest the phanerogamist, it produced many important studies on fossil angiosperms. Among these the earlier works of Berry in America (e.g., 1911) and the studies of the Reids in England (e.g., 1908) may be specially noted, the one adding to our knowledge at the earlier end of the angiosperm time scale, the other at the later end. Second, there was a great development in the study of glaciation and its possible consequences, particularly as regards the Pleistocene. It may be claimed that our modern conceptions on this subject date from these years, which saw the publication of Penck and Brlickner's Bie Alpen in Eiszeit- alter (1901-1909) , as well as much of the work of Andersson, de Geer, and others. This is perhaps the most appropriate point also for a brief reference to the study of the distribution of cryptogamic plant groups. Because the spermato- phytes reproduce by what are usually macroscopic seeds, because they generally have bulky and resistant plant bodies, and because they have a relatively short geological history, their plant geography has particular values of its own which must not be applied to other groups, though each group has its own relation to this subject. Unfortunately, for the lower plants, much less of the necessary background knowledge is available and the practical difficulties are greater; thus it is fair to say that the only groups of cryptogams which have received the same kind of geographical treatment as the seed plants are the easily collected groups of the ferns and mosses. Christ's standard work on ferns, the Geographie der Fame (1910) dates from these prewar years, but the main work of this period, Herzog's Geographie der Moose, was rather later (1926). Most groups have of course received incidental treatment in the course of taxonomic studies, as for instance in the two editions of the Pflanzenfamilien, but, apart from those mentioned, the only plants which need comment are the lichens, another easily collected group, some of which have long attracted attention by reason of their extraordinarily wide ranges. Indeed, crytogamic phytogeography is largely an untilled field, but it is also one with special difficulties of its own, chiefly inherent in the much longer and more hazy geological history of these plants. About the time of the first World War another switch of emphasis became apparent. As already explained, Schimper had long since pointed out that exist- ing floras exhibit only one moment in the history of the earth's vegetation and that in consequence the history of the earth's surface is a matter which must deeply concern the plant geographer. About 1915 several circumstances con- spired to focus attention on this aspect of the subject, so that what had in fact always been its core became crystallized more definitely than hitherto into what has now become known as historical plant geography. Among these cir- 756 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES cumstances was certainly the normal swing of the pendulum, in this ease from the extreme of formal taxonomy and purely descriptive ecology. Two other not clearly related influences, however, were more particularly concerned. These were the publication by Willis (1922) of his theory of age and area and the publication by "Wegener (1924) of his theory of continental displacement. There is much evidence for the belief that the success of a scientific theory has often been due to the fact that a general combination of trends and circum- stances, not in themselves easily discernible, have served to predispose public opinion favorably toward it, almost as if an unconscious sort of propaganda had been at work. There is little doubt, for instance, that this is broadly true of Darwinism itself, which, when the time was fully ripe, became widely estab- lished with what was really remarkable rapidity and unanimity. So with the two theories just mentioned. They were something new when the times were set for novelty and because of this, and also perhaps because each contained an element of the mysterious, they gained considerable attention. Willis' work appeared straightforwardly as a contribution to botanical thought, but it developed from an evolutionary approach to the subject. Readers may find elsewhere (e.g., New Phytol. [1951] , 50 : 135) accounts of it longer than is appropriate here. It need only be said that Willis was attracted to the subject of plant geography mainly because of the way in which it seemed to him capable of helping toward a better understanding of the processes of organic evolution and particularly because of the way it might be made to afford evidence against the theory of natural selection, which Willis' experience caused him to criticize. Very briefly, Willis maintained that the choice lay between natural selection and mutation and that, since mutation requires no assumption of a widespread supersession and elimination of "unfit" species, which is inherent in the con- ception of natural selection, any detection of an exponential rate of speciation and spread might be held to indicate that mutation rather than selection had been the paramount process in evolution. Such an exponential rate Willis claimed to demonstrate in the "hollow curve" type of graph. As a projection of this, as it were, Willis argued that under continuous mutation, not only would the totality of species constantly increase, but, barring accidents, the longer a species existed the wider would be its spread, and it is by this conception of "age and area," as he called it, that his scientific work is most familiar. This is not the place to attempt an appraisal of Willis' theories or an assess- ment of his direct contribution to plant geography as such. His claim to a place in the history of that subject is based on something rather different, for the service that he rendered was that of provoking (to use the ynot juste) a renewed interest in the whole science. But this does not altogether explain the almost violent reaction that many of his opinions occasioned and there were, it seems, two other reasons. One certainly was that he had the temerity to question the long popular theory of natural selection; the other, that he puts into words what many felt. For both these different reasons his writings received a measure of publicity and criticism which possibly surprised no one more than their aiithor. As we now look back through the years, the nature of Willis' achievement in plant geography has become clearer. It is that he showed, even if without first intent, that plant geography was not the exhausted subject which had yielded place to more modern disciplines but was a living one which still posed pro- GOOD: PMNr GEOGRAPHY 757 found problems of fundamental importance. In short, he did much to restore to it the prestige which, during the previous generation, it had seemed to lose; and it is important to realize this because it helps to relate his work to that which we must now go on to notice. Wegener's theory of continental displacement, or "drift" as it is sometimes called, dates from about 1915, though it did not become common currency until after the war, and its basic conception was not altogether novel. The relation between ideas of continental displacement and plant geography is a double one. First, the facts of discontinuity or disjunction, using those terms in their widest sense, clearly provide some circumstantial evidence for or against the view that displacement may have occurred; second, the idea is valuable to those who would explain the fioristic relationships which today exist between the sepa- rate continental masses. Both aspects combine to make drift a particularly live problem for plant geographers. Up to the present, and despite an enormous amount of study from various points of view, ideas of continental movement remain hypothetical. No summing up of the matter is really possible, but the situation in relation to the geography of the flowering plants, as it stands today, can be stated quite shortly. For at least one hundred years — which means in effect ever since the subject of plant geography took real form — there has been a common belief that the leading facts cannot be explained at all satisfactorily so long as it is held that the geography of the world, and particularly the isolation of the continents, has always been as it is now. In this connection it is well to remember that there are two ways in which a junction may be effected between separated entities. One is by inter- posing something in such manner as to bridge the gap between them; the other is by moving one or other of them bodily until the two come into contact. The first method, since it did no violence to generally accepted beliefs, was for long the accepted explanation; but it has aspects which, purely from the point of view of the plant geographer, make it a less attractive proposition than the second. As for the more direct geological and other evidences for displacement, these are at present generally held to be inadequate, and those who favor this theory are therefore faced with the fact that, on this ground, it is not generally acceptable to geologists and geodesists. How far this essentially negative attitude of objec- tion is justified time alone will show; but there are not a few who feel that the rejection of a hypothesis simply on the ground of inexplicability is unwise. Finally, with regard to Wegener's theory it must not be forgotten that it in- volved not only continental displacement, but also the idea of a more or less continuous movement of the poles. Any such movement would of course in turn involve corresponding movements in the climatic zones of the world, and a pos- sibility of this kind as an explanation of many difficult phytogeographical facts has scarcely been sufficiently examined as yet. Early in the interwar years there came two important developments which derived directly from the earlier work on glaciation already mentioned. The more important was the growth, under the leadership of Erdtman (1943) and others, of the technique of pollen analysis. This technique made it possible to form various postulates from the proportionate occurrence of different kinds of pollen grains in peat and similar deposits about the nature of the vegetation contemporary with the deposits and thus to draw a much more complete picture 758 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES of the general conditions of the time. Strictly, pollen analysis is on the borders of plant geography, especially where that subject impinges on archaeology, but within its limitations it has been and doubtless will continue to be a valuable adjunct. The second development was the discussion which grew up around what is usually called the "nunatak theory," the view that some elements of the pre- glacial floras survived those ages on refuges actually within the ice-cap but not themselves glaciated. This idea again was not in itself new, but in the nineteen- twenties it received much fresh impetus from the explorations of Fernald in the region of the St. Lawrence River, and from his energetic writings (e.g., 1925). The general suggestion of which this was a particular expression is an attractive one, namely, that tlie explanation of certain puzzling phytogeographical facts is to be found in the occurrence of refuges where plants may have been able to avoid the worst consequences of climatic change and survive. But there is as yet no overwhelming evidence that it is necessary to invoke this explanation. The last few paragraphs deal with matters which, despite their apparent diversity, nevertheless have a considerable common element, revealing clearly the main trend in the study of plant geography between the two wars, namely, its concentration on the historical-developmental aspects of the subject. Major works in this tradition soon appeared, and price of place may be given to Irmscher's Pflanzenverhreitung und Entivicklung der Kontinente, published in 1922 and followed in 1926 by Hayek's AUgemeine Pflanzengeographie, which, although much more of a textbook, had the same approach. Both of these are important, but it is fair to regard a later book as the real primer of the new interest. This was Wulff's An Introduction to Historical Plant Geography (1943), which was composed much more in the new idiom than either of the earlier books. This publication is important, too, as marking the entry into the field of the great new Russian school of botanists which had grown up since the Revolution, a school whose full influence is still impeded by barriers of alphabet and language. Fortunately Wulff's first volume, which was published in 1932, was translated into English during the war, but his second and much larger volume is still available only in Russian. This is also true of the monumental Flora URSS edited by Komarov, begun in 1934 and still in active progress. It contains a great mass of information about the distribution of plants over the huge but hitherto little studied tracts of much of central and eastern Asia. This is perhaps the best place at which, ignoring chronology for the moment, to refer briefly to two later publications, because, with that of Wulff, just men- tioned, they form a mutually complementary trilogy, covering with reasonable adequacy most aspects of modern plant geography and giving as complete a picture of the present situation as can, in all circumstances, be expected. These are Cain's Foundations of Plant Geography, which appeared in 1944, and the present writer's book, The Geography of the Flowering Plants, written before the late war but unavoidably delayed in production until 1947. Both have his- torical plant geography as their chief emphasis, but while the former is of spe- cial interest for its treatment of many particular aspects of the relation between evolution and plant geography, such as polyploidy, the latter is rather more a review of the facts of angiosperm distribution and a reconsideration, from a developmental point of view, of the factors which have caused them. All three GOOD: PLANT GEOGRAPHY 759 of these books take, at least as a partial basis, the theory of tolerance, published in 1931, by the author of the last of them in an attempt to integrate into some generally applicable working hypothesis of plant geography the many "factors of distribution." Returning now to a more general consideration of the interwar years we find that the literature of plant geography is so extensive that it is difficult to select from it, though several broad features demand comment. One of these is the special attention given, notably by German workers, to the more detailed study and analysis of types of distribution area, or areography as it has come to be called, a subject to which Hannig and Winkler's new serial publication Die Pflanzenareale, founded in 1926, contributed much and of which there have been various minor reflections in more recent years. Another noteworthy and valuable feature of this period was the large number of memoirs written about various phytogeographically strategic parts of the world by authors fully conversant with their floras. Anything like an exhaustive list would be much too long here, but they may be exemplified by the work of several authors : Allan and Oliver for New Zealand; Bews for South Africa; Perrier de la Batliie for Madagascar; Gleason for the mountains of southern Venezuela; Guillaumin and others for New Caledonia; Setchell and many others for various parts of the Pacific; Scotts- berg and St. John for Hawaii; Skottsberg for Juan Fernandez; Lam and van Steenis for parts of Malaysia; Merrill for the Philippines, and Hulten for the region of the Bering Strait. With all this went many advances in cognate subjects, as, for instance, the study of angiosperm fossil floras in which the work of Berry and, rather later, Chancy was notable, but for the rest it must suffice to mention, as representative of many others, three books which, though very different from one another, nevertheless each added something of worth to the general store. The first of these, in order of appearance, was Marcel Hardy's little book The Geography of Plants (1925), the uninspired title of which may well have served to obscure its very real merit as one of the few really concise and readable general accounts of world vegetation as a whole. The second is the volume of essays published in honor of W. A. Setchell (Goodspeed, ed., 1930) in which a number of eminent phytogeographers give authoritative accounts of their own special interests. The third is Ridley's great book The Dispersal of Plants Throughout the World (1930), in which there is surely gathered together all that is known, or at least was known at that time, about this peculiarly bewildering subject. Probably no other branch of plant geography has been so misunderstood, and even so much misrepresented, as this, and Ridley, although he confines himself largely to the recording of facts, at least provides some kind of sheet anchor, so that the subject may be more safely approached. Of the years since the outbreak of the second World War not much can be said here. They are too close to allow us to generalize and it can only be said that the literature is still copious and shows no sign of abatement. One or two items of this period have already been referred to and to these a few others must be added. Two publications of the actual war years testify to the continued vitality of what has been called here the German school, namely, three papers by Vester (e.g., 1940), in which he summarizes, with the help of small maps, the distributions of all the families of angiosperms, and Meusel's book Vergleichende 760 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES Arealkunde (1943), which is in many respects a textbook of areography. More recent is the great Dutch project of the Flora Malesiana (van Steenis, ed., 1948 — ), which, altliough in the main taxonomic, is so broadly based on geo- graphical principles that it can scarcely fail to add enormously to our knowledge of the distribution of plants in a particularly significant part of the world. There are also numerous accounts of the phytogeography of sundry parts of the north- ern temperate regions and to represent these there can be no better choice than Hulten's beautifully produced Atlas (1950), which deals exhaustively with the geography of the Scandinavian flora. Of very different scope is Willis' third and largest book The Birth and Spread of Plants (1949) which, somewhat hidden by a rather awkward kind of presentation, contains much of real importance. The war itself was responsible for not a little progress in the paths of plant geography. The extension of hostilities into the great and relatively unfamiliar spaces of the Pacific, especially, caused a considerable revision and augmentation of our knowledge of those regions, in which the botanists took their fair share, as is instanced by Merrill's useful and delightful account (1946) of tropical vegetation in that area. But the over-all impression of these later years, and one indeed hopes that it is a true estimate, is that there is coming about a much needed reintegration of the different branches of plant geography. This may be illustrated by reference to three points. First there is the growing tendency for purely taxonomic works, such as floras and systematic monographs, to pay more attention to the geography of their subjects, to bring together as much geographical information as they are able to and often to illustrate it with maps. This is desirable enough in itself, but it is also an enormous help to the phytogeographer, who must rely so much on reliable taxonomy for the facts which he endeavors to interpret. Second, one has only to look at recent volumes of the periodicals devoted to ecology to see how much the horizons of the ecologists have widened and how much more concerned with the main stream of plant geography they are becoming. Finally, there is the development which, more than anything else perhaps, characterizes the postwar years, namely, the growth of that combination of taxonomy, geogra- phy, and genetics which has come to be known as cytogeography, itself so essentially a synthetic effect. It is very apposite, in this volume which celebrates the first hundred years of the California Academy of Sciences, to mention a publication which best typifies this latest phase in the development of plant geography, Babcock's great study of the genus Crepis, which appeared in 1947. It is further work of this kind, based on phytogeographically significant plant groups, that is more likely than almost anything else to speed the progress of plant geography. At the end of the first century of the Academy's history, what are the chief impressions 1 Two seem particularly noteworthy. One is the truth of the aphorism, Plus ga change, plus c'est le meme chose. Probably no previous hundred years has seen such profound changes as this latest century, certainly not in the scien- tific world, and yet one cannot help feeling that, if Alphonse de Candolle and some of the other pioneers of plant geography could once more walk the earth, they would understand us and our problems pretty well, though we might for a time speak in rather different dialects. Advance in knowledge since their time has indeed been enormous, but it has been am])lification rather than violent GOOD: PLANT GEOGRAPHY 761 chanD:e, evolution rather than revolution. There have been secessions, but there have also been federations, and the main outlines of the subject now are very much what they were a hundred years ago. The difference is that we have a deeper understanding of them. A second impression, or so it seems to the present writer, is that today, just as in 1853, we stand on the threshold of great advances in biological thought and method. Our particular subject, plant geography, involves the former rather than the latter, but the indications of future change are not far to seek. There is an increasing impatience with ideas which owe their perpetuation more to tradition than to logic, and for many there is a growing doubt of our ability to arrive at the answers of some of our most urgent problems with our present major postulates. It is easy enough, one realizes, to forecast change when there is no obligation to foretell its shape, but its prognostication may at least help us to be ready for it and to take advantage of it. To this end there are two aids. The first is austerity, or perhaps asceticism is a better word, in scientific thought and theory. Today the pressure of events and many other influences combine to make more than ever difficult the pursuit of truth for its own sake, and it must not be forgotten that this is the only real road to scientific progress. There is need too for a higher standard of logical argument and a stronger guard against facile generalizations and false conclu- sions. The second requisite is receptiveness and suppleness of mind which, it is perhaps worth stressing, is in no way antipathetic to intellectual integrity. Every new idea, however fantastic it may appear at first sight, is entitled to critical consideration, and the wise man will treat none with complete contempt. It has been well said that all great truths begin as heresies and that all new knowledge contradicts the old. However true this dictum may or may not be, it is at any rate an admirable motto for the study wall of the plant geographer. BIBLIOGRAPHY Babcock, E. B. 1947. The Genus Crepis. Pt. I. The Taxonomy, Phylogeny, Distribution, and Evolution of Crepis. Univ. Calif. Publ. Bot., 21. xii + 197 pp. Berkeley and Los Angeles. Bentham, G., and J. D. Hooker 1862-1883. Genera Plantarum. 1(1862-1867); 2(1873-1876); 3(1883). London: Reeve and Co. Berry, E. W. 1911. The Lower Cretaceous Flora of the World. In Maryland Geol. Surv., 1896 — Cretaceous . . . 1911, pp. 99-151. Cain, S. A. 1944. Foundations of plant geography. 556 pp. New York: Harper. Candolle, a. L. P. P. DE 1855. Geographie Botanique Raisonnee. Vol. I, xxxii + 606 pp.; Vol. II, 607-1365 pp. Paris: V. Masson. Candolle, A. P. and A. L. P. P. de 1824-1873. Prodomus Systematis Naturalis Regni Vegetabilis. 17 vols. Paris: Treutel et Wiirtz. 762 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Christ, H. 187.9. Das Pflanzenleben der Schweiz. xiv + 488 pp. Zurich: F. Schulthess. 1910. Die Geographie der Fame. 357 pp. Jena: G. Fisclier. Clements, F. E. 1916. Plant Succession. Carnegie Inst. Wash. Publ. No. 242. 512 pp. Washington. COWLES, H. C. 1899. Tlie ecological relations of the vegetation on the sand dunes of Lake Michigan. Bot. Gaz., 27:95-116, 167-202, 281-308, 361-391. Darwin, Charles 1859. The Origin of Species by Means of Natural Selection. 6th ed., 1881. 434 pp. London: Watts. Drude, O. 1887. Atlas der Pflanzenverbreitung. 6 pp., 8 maps. Gotha: J. Perthes. 1890a. Handbuch der Pflanzengeographie. 582 pp. Stuttgart: J. Engelhorn. 1890b. Vegetationsformationen erlautert an der centraleuropaischen Flora. Bot. Jahrb. Engler, 11:21-51. Engler, a. 1879-1882. Versuch einer Entwickelungsgeschichte der Pflanzenwelt. I, 202 pp.; II, 347 pp. Leipzig: Engelmann. 1899. Die Entwickelung der Pflanzengeographie in den letzten hundert Jahren und weitere Aufgaben derselben. In Wiss. Beitr. z. Gedachtniss der hundert- jahrigen Wiederkehr des Antritts v. A. v. Humboldt's Reise nach Amerika, 1799. Ges. Erdkunde. 247 pp. Berlin: W. H. Kuhl. 1900- (Ed.) Das Pflanzenreich. 105 vols. Leipzig: W. Engelmann. 1905. Ueber floristische Verwandschaft zv^^ischen dem tropischen Afrika und Amer- ika, sowie iiber die Annahme eines versunkenen brasilianisch-Athiopischen Continents. Sitzungsber. K. Preuss. Akad. Wissen. Berlin, 1905(6) : 180-231. Engler, A., and 0. Drude (eds.) 1896-1928. Die Vegetation der Erde. 1st and 2d eds. 15 vols. Leipzig: W. Engelmann. Engler, A., and L. Diels 1936. A. Engler's Syllabus der Pflanzenfamilien. 11th ed. xiii + 419 pp., 476 figs. Berlin: Gebr. Borntraeger. Engler, A., and K. Prantl 1889-1924. Die naturlichen Pflanzenfamilien. 1st ed., 1889-1915; 2d ed., 1924- . Leipzig: W. Engelmann. Erutman, G. 1943. An Introduction to Pollen Analysis. 239 pp. Waltham, Mass.: Chronica Botanica. Fernald, M. L. 1925. Persistence of plants in unglaciated areas of boreal America. Amer. Acad. Arts and Sci. Mem., 15:241-342. Good, R. 1931. A theory of plant geography. New Phytol., 30:149-171. 1947. The Geography of the Flowering Plants. 403 pp. London and New York: Longmans, Green. Goodspeed, T. H. (ed.) 1936. Essays in Geobotany in Honor of William Albert Setchell. xxvi + 320 pp. Berkeley: Univ. Calif. Press. Graebner, p. 1895. Studien iiber die norddeutsche Heide. Bot. Jahrb. Engler, 20:500-654. GOOD; PUNT GEOGRAPHY 763 Gray, A. 1859. Diagnostic characters of new species of phaenogamous plants, collected in Japan by Charles Wright, with observations upon the relationship of Jap- anese flora to that of North America, and other parts of the north temperate zone. Amer. Acad. Arts and Sci. Mem., n.s., 6:377-452. Grisehach, a. H. R. 1884-1885. Die Vegetation der Erde nach ihrer klimatischen Anordnung. 2d ed. Vol. 1, 567 pp.; Vol. 2, 594 pp. Leipzig: W. Engelmann. GUPPY, H. B. 1903-1906. Observations of a Naturalist in the Pacific between 1896 and 1899. Vol. 1 (1903), Vanua Levu, Fiji, xix + 392 pp.; Vol. 2, Plant Dispersal, xxvi + 627 pp., 5 pis., 5 maps. London and New York: Macmillan. 1912. Studies in Seeds and Fruits. 528 pp. London: Williams and Norgate. 1917. Plants, Seeds and Currents in the West Indies and the Azores. 531 pp. Loudon: Williams and Norgate. 1917-1920. Plant distribution from the standpoint of an idealist. Journ. Linn. Soc. Bot., London, 44:439-471. 1919. The island and the continent. Journ. Ecol., 7:1-4. Hannig, E., and H. Winkler (eds.) 1926- Die Pflanzenareale. 5 vols.. Vol. 2 with L. Diels; Vol. 3 with G. Samuelsson; Vol. 5, maps. Jena: G. Fischer. Hardy, M. 1920. The Geography of Plants. 327 pp. Oxford: Clarendon Press. Harshberger, J. W. 1911. Phytogeographical Survey of North America. Ixiii + 790 pp. New York: Stechert and Company. Hayek, A. 1926. Allgemeine Pflanzengeographie. 409 pp. Berlin: Gebr. Borntraeger. Herzog, T. 1926. Geographic der Moose. 439 pp. Jena: G. Fischer. Hitchcock, A. S. 1898. Oecological plant geography of Kansas. Acad. Sci. St. Louis. Trans., 8:55-69. Hooker, J. D. 1853. Introductory Essay to the Flora of New Zealand. London. 1859. On the Flora of Australia, its Origin, Affinities and Distribution; being an Introductory Essay to the Flora of Tasmania. 128 pp. London: L. Reeve. 1875-1897. The Flora of British India. 7 vols., in 24 parts (1872-1897). London: Reeve and Co. Hulten, E. 1950. Atlas over Vaxternas Utbredning i Fanerogamer och Ormbeinksvaxter Norden. 512 pp. Stockholm: Generalstabens. Humboldt, A. von 1817. De Distributione Geographica Plantarum, Secundum Coeli Temperiem et Altitudinem Montium Prolegomena. 249 pp. Paris. Irmscher, E. 1922. Pflanzenverbreitung und Entwickelung der Kontinente. Studien zur genetischen Pflanzengeographie. Hamburg Inst. Allgem. Bot. Mitteil., 5:17-235. 764 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Karsten, G., and H. Schenk (eds.) 1904-1944. Vegetationsbilder. 26 vols. Jena: G. Fischer. KoMAROv, V. L. (ed.) 1934-1952. Flora URSS. 18 vols. Moscow and Leningrad: The URSS Acad. Sci. Press. 1941. Materials on the History of the Flora and Vegetation of the URSS. Fasc. I. 413 pp. Moscow and Leningrad: The URSS Acad. Sci. Press. (English sum- maries of Russian papers.) Krasnov, a. 1886. Geobotanical researches in the Kalmuk steppe. Russ. Geog. Soc. St. Petersb. Bull., 22:1-52. Merrill, E. D. 1946. Plant Life of the Pacific World. 295 pp. New York: Macmillan. Meusel, H. 1943. Vergleichende Arealkunde. 2 vols., Vol. 1, 437 pp. Berlin: Gebr. Borntraeger. Penck, a., and E. Bruckner 1901-1909. Die Alpen in Eiszeitalter. 3 vols. Leipzig: C. H. Tauchnitz. Pound, R., and F. E. Clements 1900. The Phytogeography of Nebraska. 442 pp. 2d ed. University of Nebraska. Raunkiar, C. 1907. Planterigets Livsformer og deres Betydning for Geograflen. 132 pp. Copenhagen. Reid, C, and E. M. Reid 1908. On the pre-glacial flora of Britain. Journ. Linn. Soc. Bot., London, 38:206-227. Ridley. H. N. 1930. The Dispersal of Plants throughout the World. 744 pp. Ashford, England. RUBEL, E. 1927. Ecology, plant geography and geobotany, their history and aim. Bot. Gaz., 84:428-439. 1930. Pflanzengesellschaften der Erde. 464 pp. Bern-Berlin. Sargent, C. S. 1884. Report on the forests of North America. 612 pp. Washington: Government Printing Office. SCHIMPER, A. F. W. 1898. Pflanzen-Geographie auf physiologische Grundlage. (1903) 876 pp. (Eng. trans, by Groom and Balfour, 1903. Oxford.) Jena: G. Fischer. SCHROTER, C. 1912. Genetische Pflanzengeographie. Jena. ScHROTER, C, and J. Fruh 1904. Die Moose der Schweiz. Berne. Solms-Laubach, H. 1905. Die leitenden Gesichtspunkte einer allgemeinen Pflanzengeographie in kurzen Dorstellung. 243 pp. Leipzig: A. Felix. Steenls, C. G. G. J. van (ed.) 1950- Flora Malesiana. Djakarta: Noordhoff-Kolff. GOOD: PLANT GEOGRAPHY 765 Tansley, a. G. 1911. (Ed.) Types of British Vegetation, xx + 416 pp. Cambridge Univ. Press. 1947. Tlie early history of modern plant ecology in Britain. Journ. Ecol., 35:130-137. Thistelton-Dyer, W. T. 1893. Alphonse de Candolle. Nature, 48:269-271. Vester, H. 1940. Die Areale und Arealtypen der Angiospermen-Familien. Eot. Archiv, 41:203- 275, 92 figs. Warming, J. E. B. 1895. Plantesamfund. vii + 335 pp. Copenhagen: P. G. Philipsen. Wegener, A. L. 1924. The Origin of Continents and Oceans. (Eng. trans, from 3rd German ed. by J. G. A. Skerle.) xx + 212 pp. London: Methuen Co. Willis, J. C. 1922. Age and Area. A Study in Geographical Distribution and Origin of Species. 259 pp. Cambridge: Univ. Press. 1949. The Birth and Spread of Plants. Geneva. WULFF, E. V. 1943. An Introduction to Historical Plant Geography. (Eng. trans, by E. Brissenden.) 223 pp. Waltham, Mass., U. S. A. ANIMAL GEOGRAPHY By KARL P. SCHMIDT Chicago 'Natural History Museum The comprehensive work on the subject of Animal Geography, published in 1853 by Ludwig Schmarda, of the University of Gratz, serves very well as a summary of the state of knowledge in this field in the eighteen-fifties. Die geo- graphische Verhreitung der Thiers devotes 93 pages (with no less than 129 pages of notes and references) to the modality and causality of animal distribution, in which he discusses the influences of heat, light, air, electricity, climate, seasonal cycles, and food, much as the ecological factors in animal distribution are set forth today. In this section it is apparent that the data were in every respect inadequate for a comprehensive review in 1850. Schmarda goes on to discuss the dependence of animals on their medium and substrate, the altitude distribu- tion of both land and marine animals, general ideas about dispersal, and the concepts of faunas and zoological regions. He has taken the step, bold enough for 1853, of giving up the concept of a single center of creation and of dispersal in favor of a number of such centers. He is aware also of the phenomenon of vicariation, of the replacement of one species of animal by an obviously related one in adjacent areas or regions. His twenty-one terrestrial and ten marine regions are rather casually chosen and as casually characterized by some dominant group — the Middle European Realm, for example, is the realm of insectivores and of carabid and staphylinid beetles, IMadagascar the realm of lemurs. The discussion of the terrestrial regions occupies 143 pages, with 258 pages of references, and that of the marine regions 58 pages with 94 pages of notes. A colored Mercator map delimits the 31 realms. On the eve of the revolution in zoological thought brought about by Darwin's Origin of Species, the delimitation of the principal faunal regions of the world was the principal preoccupation of zoologists interested in distribution. Thus in 1858 P. L. Sclater set forth the principal terrestrial regions as indicated by birds, and in the same year Albert Glinthor did the same for reptiles, with a very fair agreement between the two distinct approaches. The appearance of Darwin's Origin of Species in 1859 marked a radical change of direction and an enormous stimulus to botanical and zoological explora- tions and studies in every field. By Darwin's time, the broad patterns of animal distribution shown by the larger and otherwise more conspicuous animal types had been made known, and the existence of these patterns presented increasing diffi- culties to theories of special creation. Thus Darwin's summary of the evidences from animal distribution that favored the evolutionary origin of animal types (whether species, genera, or higher groups) marked the end of the purely descrip- tive era, and the beginning of a period of interpretation and speculation and re- examination of the phenomena in the field of geographic distribution of plant and animal life, as in all other segments of biology. [767] 768 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES .«U^ 4 ^ ^ >«4>wgw>«gi^ *gKgggg'lljiliaiMWglt5WgW i|iaMwaw -S n. t' i^ u 3 \ VY- Kit /r' " f \ } \ -i ■ Sr-"*^ -[■■ ■ 4fH .'/ £S / X - f f - ■ it ■■■ 4f X. Xj « Figure 1. Schmarda's attempt to divide the world into zoological regions (see opposite page). From Schmarda, 1853. SCHMIDT: ANIMAL GEOGRAPHY 769 Darwin had been strongly influenced in his whole evolutionary trend by the facts of animal and plant distribution. He devotes two chapters in the Origin of Species to this subject, and in these he discusses the genetic similarities within the faunas and floras of the several continents; the means of dispersal available to plants and animals; the influence of glacial periods on distribution; the distri- bution of freshwater animals; and the inhabitants of oceanic islands. In illustra- tion of the last mentioned topic he deals especially with the animals and plants of the Galapagos Islands, where the phenomena of insular distribution had so greatly impressed him in 1835. Alfred Russell Wallace, Darwin's friend and fellow evolutionist, had lived for many years in tropical regions, first in the Amazon Basin and later in the East Indies, where he had been especially impressed by the phenomena of animal distribution. He thus had a broader and more direct and intimate acquaintance with the subject than any other naturalist traveler of his century. He was con- tinually at work on this subject from 1860 until 1876, the date of publication of his two volumes on The Geographical Distribution of Animals. He somewhat modestly refers to this work as an extension and amplification of the two chapters on the subject in the Origin of Species, comparing it with Darwin's own two- volume expansion of the chapters on animals and plants under domestication. The two principal sections of Wallace's work are contrasted as "zoological geog- raphy," a descriptive discussion of the land animals of the different zoogeographic regions, and "geographical zoology," a review of the distribution of vertebrates and certain invertebrates, group by group. This work was ponderous and at the same time naive in supposing that "a solution of zoogeographic problems could be attempted with some prospect of success." It remains a necessary work to a specialist in this subject, but modern changes in classification, the great advances in paleontology, and the rise of ecology, combine to make it useless as an intro- duction to animal geography. The more popularly written Island Life (1880) is Figure 1. General map of the geographic distribution of animals, drawn up by Ludwig K. Schmarda, 1852. The land realms are: I. the Arctic, realm of fur bearers and aquatic birds; II. Central Europe, realm of insectivores and staphilinid and carabid beetles; III. Caspian Steppe, realm of the saiga antelope and of burrowing rodents; IV. Central Asian Steppes, realm of the horse tribe; V. European Mediterranean, realm of heteromeran beetles; VI. China, realm of pheasants; VII. Japan, realm of the giant salamander; VIII. North America, realm of rodents, conirostrine and dentirostrine birds; IX. Sahara Desert, realm of the ostrich and of melasomas (tenebrionids) ; X. West Africa, realm of catarrhine monkeys and termites ; XI. Highland Africa, realm of ruminants and pachy- derms; XII. Madagascar, realm of lemurs; XIII. India, realm of carnivores and pigeons; XIV, the Sunda World, realm of snakes and bats; XV. Australia, realm of the marsupials, monotremes and meliphagid birds; XVI. Central America, realm of land crabs; XVII. Brazil, realm of edentates and platyrhine monkeys; XVIII. Andean, realm of the llama and condor; XIX. Pampa, realm of the viscacha and "harpalid" beetles; XX. Patagonia, realm of Darwin's rhea and of the guanaco; XXI: Polynesia, realm of nymphalid butter- flies and of the Apteryx (New Zealand). The marine realms are: XXII. Arctic Ocean, realm of marine mammals and amphipod crustaceans; XXIII. Antarctic Ocean, realm of marine mammals and penguins; XXIV. North Atlantic Ocean, realm of cods and herrings; XXV. South European Mediterranean, realm of the labrid fishes; XXVI. North Pacific Ocean, realm of the scombrid and mail-cheeked fishes; XXVII, Tropical Atlantic Ocean, realm of plectognath fishes, manatees, and of pteropods; XXVIII, Indian Ocean, realm of buccinoids and hydroids; XXIX. Tropical Pacific Ocean, realm of corals and holothurians; XXX. South Atlantic Ocean; XXXI. South Pacific Ocean. 770 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES 'fc'-^ 1^3 II r* 1 if-W ^^; e^iy ""^i • j = i ! >y t , ' ^ ■ ■: I- :* I- '"■'.. ^ -9- - ■ ' '^ ''>. /A -Ami i '-J- J ,- . ^A^^fi^ Figure 2. Wallace's zoogeographic regions. From Wallace, 1876. SCHMIDT: ANIMAL GEOGRAPHY 771 Wallace at his best. Freed of much of the cumbrous and now obsolete detail that burdens the larger work, the later volume remains a satisfactory and vivid introduction to the study of animal distribution. Unfortunately the long and detailed discussion of the causes of glacial periods rather arbitrarily inserted in this work seems to a modern or skeptical reader incredibly optimistic in its over- simplification of so complex a subject. A completely revised edition of Island Life, retaining as much as possible of Wallace's own writing, is much to be desired. In broad outline, the historical development of zoogeography is dominated by Wallace in the generation between the eighteen-fifties and the date of Island Life. This generation concerned itself with an accumulation of the facts of dis- tribution in what might be called "Descriptive Animal Geography"; and with fundamental improvements in the classification of animals. The delimitation and classification of the geographic subdivisions of the earth's surface that seemed to accord best with the facts of animal distribution was perhaps the principal subject of controversy in this period, much of it based on inadequate studies or fields too limited to afford contributions of permanent value. Wallace had adopted the system of regions proposed by Sclater with only a few changes of names. Thomas H. Huxley's entry into the field in a well-reasoned paper on a limited group of birds (1868) must be mentioned, since it suggests an important reclassification and hierarchial arrangement of the zoological regions into three principal realms. An anonymous writer proposed acceptable terms for these — "Arctogaea," to include the Palearctic, Nearctic, Oriental, and Ethiopian regions; "Neogaea" for the Neotropical region; and "Notogaea" for the Aus- tralian. This, with the combination of the Nearctic and Palearctic regions into the Ilolarctic by Heilprin in 1878, leads directly to the modern grouping of realms, regions, subregions, and provinces. For all the enthusiasm of Wallace, and in spite of the fact that he and his contemporaries quite correctly assessed the revolutionary importance of the theory of evolution to the concepts of animal geography, the era in which his Geographic Distribution of Animals was the leading work in the field remained essentially involved in these static problems. There was a long series of papers involving the problem of combining the Nearctic and Palearctic regions into an over-all Holarctic region; and whether a distinct Sonoran region should be cut off from the Holarctic to include the southern half of North America. The controversy as to where to draw the line between the Oriental region and the Australian, and especially as to the place of the fauna of Celebes in this scheme, became a zoogeographic classic, which has had renewed attention in the decade of the nineteen-fortics. There was endless report and argument about the degree of relation between one faunal province and another based entirely on the existing faunas, without reference to their origin and history. Permanence op Continents Darwin and Wallace, with the American geologist Dana to support them, regarded the continents as stable features of the earth's surface, and postulated connections between the continents and between continents and islands only where they exist now, as at Panama; or where there are shallow seas, within 772 ^ CENTURY OF PROGRESS /N THE NATURAL SCIENCES the limits of the continental platforms, whieh drop off quite abruptly to the abyssal ocean floor at about the 200-meter depth line. Wallace accordingly framed the classification of islands into "oceanic" and "continental," the oceanic islands being thought to have received all of their plant and animal life overseas, whereas the continental islands were populated overland, by direct invasion. The directly demonstrable land connections between continents, and between islands and continents, that have been available for the dispersal of land animals are few. There is the existing Central American isthmus connecting North and South America; the shallowness of Bering Sea and narrowness of Bering Strait indicate that this region afforded a broad connection of North America with Asia; and except for the man-made Suez Canal, Africa is connected with western Asia, and was much more broadly connected in the past before the block-faulting that produced the Red Sea. The past connection of the British Isles with the Euro- pean mainland and of the Greater Sunda Islands (Borneo, Sumatra, and Java) with southeastern Asia are undoubted, documented physiographically by the drowned river valleys in the neighboring shallow seas. Land Bridge Speculations With these known connections to explain, the existence of animal life in certain islands and to explain the resemblances and relations between animals of one continent and another, it was perhaps natural to turn to hypotheses of other transoceanic land connections to explain such facts as the predominance of marsupials in Australia and South America, or of certain types of freshwater fishes in Africa and South America. The trend to bold hypotheses of this kind begins with the English naturalist Edward Forbes, as early as 1846. Forbes analyzed the fauna of the British Isles as to its various components, and found an element in the south of England and Ireland related principally to the life of Spain and Portugal. To explain this relation, he supposed the former (but quite recent) existence of a continental land mass projecting far out into the Atlantic. At first thought, since it was known that the sea had widely trans- gressed most continental areas for long periods in the past, it seemed logical enough that land areas might equally as well have been present where the oceans now lie. Hypothetical continents or isthmuses — "land bridges" — between conti- nents were proposed and drawn in upon maps, and presently received names. An "Atlantis" was thought to have occupied most of the North Atlantic, and an "Archhelenis" the South Atlantic. Antarctica was renamed as "Archinotis." "Lemuria," constructed to account for the distribution of lemurs, spread across the Indian Ocean from India and Ceylon to Madagascar. "Pacelia" was a lobe of hypothetical land connecting the Hawaiian Islands with North America. Such speculations were strongly re-enforced by the geological hypothesis of a "Gondwana Land" uniting all of the southern continents in Paleozoic and much of the Mesozoic time introduced in 1860 by the French- American geologist Marcou for the Jurassic, modified by Neumayr (1883 and 1887), and by Suess (1885), who placed its origin in the Paleozoic. The Gondwana Land hypothesis was based on the distribution of the remarkable fossil fern Glossopteris of Permian age, and on attempts to delimit continental borders in earlier geologic ages in the light of evidence from marine fossils. The great work of Edward SCHMIDT: ANIMAL GEOGRAPHY 773 Suess, Das Antlitz der Erde (in which Gondwana Land was first so named) laid off the earth's surface in broad and bold tectonic outlines. Suess' ideas received support from the most eminent of geologists, as in Neumayr's later editions of the Erdgeschichte, Emil Ilaug's Traits de Geologie, and in Britain from J. W. Gregory. With such notable geological precedent, any specialist on any group of animals felt free to explain disconnected distributions by hypotheses of tongues of land extending over any water barrier that might separate even the species of a single genus; and such liypotheses can only be described for the era between 1880 and 1915 as "untrammeled." The bridges became ever more complicated — R. F. Scharff, for example, thought that eastern and western Australia had been connected with Antarctica by two separate land corridors. Tongues of land were thought to be of short duration, lasting just long enough for the author's purpose and not so long as to allow additional and confusing emigrations and counteremigrations to take place. During the era of land-bridge speculation eminent specialists in various fields became proponents of this or that pattern of connection between the southern continents. The great botanist Joseph Dalton Hooker had been impressed with the relations of the plants of southern South America, New Zealand, and Tas- mania in the course of his early exploring voyage with the Erebus and Terror, from 1839 to 1843. The remarkable and distinctive Araucarian pines found in southern South America and in the islands near New Zealand, and the antarctic beech NotJiofagus, found in Chile, New Zealand, and Tasmania, present to bot- anists exactly the kind of geographic relations that had led zoologists to speculate about direct land connections across existing oceans. The Swiss zoologist Riiti- meyer, in an essay on the origin of the animal life of Switzerland, published in 1867, makes the suggestion of the former existence of a vast Antarctic continent, connecting all of the southern continents and New Zealand, and this idea received support from T. H. Huxley in 1870. The Antarctic and Pacific continents then expand and contract in the minds of F. W. Hutton (1873), Theodore Gill (1875), Hermann von Ihering (1891), H. 0. Forbes (1893), Charles Hedley (1895), H. F. Osborn (1900), and A. E. Ortman (1901). All of this land-bridge history is essentially independent of the geological theories of a Gondwana Land. Ideas of land bridges were integrated with the ideas of zoogeographers by Theodor Arldt, in Die Entwicklung der Kontinente und ihrer Lehewelt in 1907 (2d ed., 1936-1938). There is a little popular sum- mary of the subject by Hans Gadow, in The Wanderings of Animals (1913). Throughout all of this era of "land-bridge building," a few zoogeographers held to the basic assumption of the permanence of the existing continents and the distinction between oceanic and continental islands. Darwin wrote skeptically about "those who make continents as easily as a cook makes pancakes." Against much polemical sniping, Wallace held firm to his original position. Georg Pfeffer, almost alone among malacologists, held out against the too easy explanations of direct and multiple land connections. Anton Handlirsch took up the polemic cudgels and showed how disgracefully superficial and how incredibly arbitrary and thoughtless had been the "creation" of land bridges. By mapping the hypo- thetical connecting land areas proposed by his colleagues, he showed that almost every bit of existing ocean bottom had been raised and lowered. As the fore- 774 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES J— -'-.r5gjt--,-r,;3g.- 2wr'7j\-»i^ift TrftB8gressiocsgebi<^t iffl Uittele^rboa. I '^&MS3 KsatiooBte im UntBrcarbGo. ^^^^ Traiisg»sii9B«geM»t Un ob«nt«it OiHwu. f Si-":" " &f .' " ,Sr'' " tlTf"'"' W~' "ii?" Sontinente wibread der filteren Tertiflrzeit. KaBtiit!»it» iiB QiigooSn (nsoii Kskfa), Oiigswtos TtMisgre«B9!Mgel))8ts, Ndjb Kut«, t'orvcls (lid IWi asa Figure 3. Land-bridges in Carboniferous and early Tertiary times, according to various authors. From Arldt, 1907. SCHMIDT: ANIMAL GEOGRAPHY 775 most authority on fossil insects, Handlirsch analyzed the existing faunas of the continents and showed that even for this ancient and eminently terrestrial group the proposed connections of continents in the southern hemisphere are flatly opposed by a vast mass of contrary evidence. Only an uncritical enthusiasm could maintain them on faunal grounds alone; as we shall now see, there is crucial geological evidence against them. ISOSTASY The whole scheme of rising and sinking continents is now found to be opposed by unshakable geological evidence, from the facts summarized as "isostasy," which show that the continental platforms are indeed stable. The whole mass of books, journal articles, and addresses to scientific meetings on the subject of land connections is an incredibly futile chapter in the history of animal geography. When the survey of the earth's surface had advanced to large-scale operations it was discovered that astronomical determinations of latitude were sharply at variance with direct measurement. The north-south breadth of Puerto Rico as found by astronomical calculation, for example, differs by about a mile from the 50-mile direct measurement. This difference results from a deflection of the plumb line toward mountain masses and toward continental masses. J. H. Pratt reported in 1855 "On the Attraction of the Himalaya Mountains and of the Elevated Regions Beyond Them Upon the Plumb Line in India." Since that time a gravity survey of the world has been made by means of the pendulum observations begun by George Biddell Airy at about the same time. The result of the world survey, which constitutes the science of geodesy, has been to re-enforce the validity of the principle of isostasy. It is found that the granitic materials of the mountains, the "sial" of Suess, have a density distinctly less than that of the basaltic rocks underlying the oceans, the "sima," in the proportion of 2.7 to 3.0. The sial, in a sense, floats on the heavier substratum of sima, and it is the sima that forms the ocean bottoms. The continental bases extend deep into the sima, with further great downward extending masses beneath the mountain ranges, except where their isostatic adjustment is not complete, as shown by the occurrence of earth- quakes. An example of continuing isostatic adjustment familiar to us in North America is the rise of the northeastern quarter of the continent after the retreat of the continental glacier; the rare earthquakes in this otherwise extremely stable area are ascribable to the readjustment of the continental block with the disap- pearance of the ice load. The conviction that the continental platforms are indeed permanent in broad outline, and that the ocean floor is of very different compo- sition, became more and more an axiom of modern geology as the extension of gravity measurements failed to find exceptions to the lightness of the continental masses relative to the ocean floors. This flatly contradicts the hypotheses of vast former continents extending across the existing oceans. Continental Drift Just when this began to be realized, an ingenious alternative was afforded by the hypothesis of continental drift elaborated by Alfred Wegener (1915), from ideas already current (Taylor, Baker). ^ Wegener, impressed by the current 1. Du Toit (1937) sketches the history of the ideas involved. 176 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Upper Carboniferaus /■" .^ Older Quaternary X X, ---»i5^ Figure 4. The arrangement of the continents in successive geological periods accord- ing to theories of continental drift. From various sources. SCHMIDT: ANIMAL GEOGRAPHY 777 belief that continental connections are necessary to explain the existing pattern of animal life on the continents, suggested that the correspondence of the con- figuration of the Atlantic coast of South America with that of Africa was the result of a drifting apart of land masses formerly joined, and rent asunder in some past age. North America was thought to have drifted away from Europe in the same way. It was found possible to fit the Australian and Antarctic con- tinents against South Africa, also, with the Indian Peninsula and Madagascar to fill the wedge-shaped gap between them, and to suppose that these land masses had drifted away from Africa to the south and east and north as the Americas had drifted to the west. The distribution of past continental glaciations and the distribution of Paleozoic coal deposits were brought into harmony with the theory of continental drift by assuming a different position of the poles during Paleozoic times. The actual break-up of the vast unified original continent is thought by "Wegener and his school to have occurred at the end of the Paleozoic. Thus the theory is of little aid to those who had explained the distributions of relatively recent groups like the birds and mammals by means of land connec- tions, or to those who invoked them for distributions of groups at the level of genera, which are demonstrably much more recent. To the more confirmed pro- ponents of continental connections, however, this led merely to the outright belief that the continents had drifted back and forth- — that they had been con- nected, separated, and reconnected. The theory of continental drift has been received with considerable skepti- cism by American geologists, many of whom had had no prior belief in any continental connections other than those existing. The confirmation of the fact that the continents are in isostatic balance merely confirmed their ideas of the continents as vastly older than the life on them. In Europe a large group espoused the Wegnerian ideas, and the literature of the subject, and finally the literature of the controversy between drift and anti-drift proponents is now vast. Much of this literature, however, still assumes as axiomatic that a past connection of the continents is necessary to explain the present distribution of land life. In the very year when Wegener proposed his theory, this assumption was shown by William Diller Matthew to be invalid. Climate and Evolution The publication of Matthew's Climate and Evolution in 1915 was an event of primary importance in the field of animal geography. This paper sums up the accumulated paleontological evidence for the Tertiary distribution of mam- mals in a masterly way; it re-enforces the belief in the general permanence of continents, which had been a cardinal point with Wallace, but which had come to be more and more widely disregarded; it reverses a long accepted criterion for the place of origin of mammalian groups especially and of other animals in broad outline; and it offers a principal cause for the long-term dispersal of species, and of faunas with their environments, that links the subject of distri- bution with the whole course of evolution, and with a reasoned interpretation of the geological record. This work, in direct contradiction in many of its well- supported statements with the contemporary literature of animal geography, could not fail to have an extraordinary influence. A considerable number of 778 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES subsequent distributional studies by students or disciples of Matthew have been based on Matthewsian principles; and the modern period in zoogeography must be dated from 1915. Matthew demonstrated that no hypotheses of trans-Pacific or trans- Atlantic connections between the continents are required to explain the existing distributions of mammals, and that the Mesozoic and Tertiary bridges between Asia and North America at Bering Strait and over the shallow Bering Sea, the Isthmus of Panama, with the perhaps still earlier bridge via the unstable island area between Australia and southeastern Asia, are the only con- tinental connections for which valid evidence exists. This is documented from Matthew's unparalleled knowledge of mammalian paleontological history. As Darwin and Wallace and Handlirsch had rightly remarked, the axiom of general stability of the continents (to the outlines of the continental shelves) is essential to any orderly and critical investigation of animal distribution. Matthew could rest renewed affirmation of this axiom on the geological evidence for isostasy. The more important contribution of Climate and Evolution is the presenta- tion of an outline on the world scale, and in the grandest historical perspective, of the dispersal of land animals throughout geological history. This rests on the synthesis by Chamberlin and Moulton of the evidence from geological history as a whole for periodic cycles of great uplift of the continents with corresponding climatic extremes, diversity, and change, alternating with base-leveling by erosion ending in widespread uniformity and amelioration of climate. Matthew suggests that these large-scale changes have dominated progressive and adaptive evolution, and that ancestral types that failed to enter the main currents of evolution have either followed the climatic changes in their dispersal, to avoid the necessity of change, or have been forced by the competition of the advancing and physio- logically more progressive types into geographically peripheral regions. The examination of existing and fossil types at the peripheries of the ranges of their groups documents this broad pattern of dispersal. The continents of the southern hemisphere extend like vast peninsulas from the larger land masses of the northern hemisphere. It is demonstrable from the paleontological record that these larger northern land masses have been the main theaters of the evolution of mammals; and the southern continents are veritable museums of relict types preserved from past ages, whose ancestors are often known to have been present at earlier times in the northern hemisphere. Centers of origin are to be expected where the more advanced forms now live rather than where the most primitive forms are found. Recognizing that the evidence from paleontology becomes progressively more obscure from the well-documented Tertiary history of the Age of Mammals through the Age of Reptiles in the Mesozoic to the origins of land animals in the Paleozoic, Matthew pleads for the interpretation of the less known by means of the better known, and doubts the existence of transoceanic connections of the southern continents even in the more remote geological periods. This is essen- tially the application of the "law of parsimony," which is basic to critical scien- tific thought. The absurdity of Arldt's determination of the probability of past trans- Atlantic and trans-Pacific land connections by means of a statistical analysis of the "votes" of zoologists becomes evident after a critical examination of Climate and Evolution. This work disposes of the supposed necessity for such SCHMIDT: ANIMAL GEOGRAPHY 779 jVif Tapirs ^nK«%$^^s^^^^^i^^^»«^^Sii^Si irr.ocerosfs Figure 5. Matthew's maps showing origin and dispersal in geological time. Upper figure, the tapirs; lower figure, the rhinoceroses. From Matthew, 1915. 780 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES continental connections with much the same finality as did the Origin of Species for the necessity of the hypothesis of special creation. "Special Creation," in 1858, would have received an overwhelming majority vote from contemporary biologists. A group of students in zoology and paleontology under William King Gregory at the American Museum of Natural History in the years 1915 to 1927 came also under the influence of W. D. Matthew, in whose personality great modesty and simplicity stood out against the background of an enormous scientific prestige. The group included Alfred Sherwood Romer, Charles Lewis Camp, and Gladwyn Kingsley Noble, with many others, and a little more indirectly Emmett Reid Dunn and myself. All of us have been involved in one way or another with the problems of animal geography, and all of us have remained disciples of Matthew. I was tempted to refer to my own commentary on Matthew ( 1943 ) as consisting of "Parerga and Paralipomena"; we have tended to look a bit askance at those "who knew not Matthew"; and it had not escaped some of our colleagues that Matthew's work had become a kind of Holy Writ to his disciples. Fortunately we have now had a strong cross-light thrown on the main thesis of Climate and Evolution by a nondisciple, P. J. Darlington, Jr., of the Museum of Comparative Zoology at Harvard University, who reviews the whole matter from the evidence of the freshwater fishes, amphibians, and reptiles. These were the groups of which Matthew had least personal knowledge. Darlington shows that Matthew's account of these groups was quite inadequate and often erroneous, and that some of their major dispersals came from the tropics instead of from the northern continents; but he ends in essential agreement with Matthew in finding the Bering Sea and Bering Strait bridge adequate to explain the dispersals between the eastern and western hemispheres. The very important evidence of the freshwater fishes, long regarded as a proof of the necessity for past direct connec- tion of Africa and South America, is now regarded by George S. Myers as explain- able by long-term round-about emigration via the Bering Bridge rather than by hypothetical trans- Atlantic connections. Myers has more particularly analyzed the freshwater fish fauna of the West Indies (1938) and of Madagascar. He shows that the fish faunas of these two great island areas cannot possibly be interpreted on the theory of connection with the adjacent continents. He rejects anylate Mesozoic land connection of Australia with southeastern Asia on account of the extreme impoverishment of the primary freshwater fish fauna of the Australian region (1951). The important evidence from fossil plants has been presented and analyzed most recently by North American paleobotanists (Chancy, 1947). So far as the Cretaceous and Tertiary history is concerned, Chaney and Axelrod (to name only two) completely reject theories of land connection other than that at Bering Sea, and their account of the Tertiary history of the floras of the northern hemisphere is in essential agreement with Matthew. One of the most striking relations between the recent faunas of South America and Africa lies in the rich, and in some respects parallel, development of the great group of worm-like lizards, the Amphisbaenidae. The group is wholly unknown in the modern fauna in eastern Asia. It is therefore especially illumi- nating that a fossil amphisbaenian of Oligocene age was discovered in Mongolia by the expedition of the American Museum of Natural History. It was described SCHMIDT: ANIMAL GEOGRAPHY 781 by Gilniore in 1043. This points again to the fact that the imperfections of the paleontological record must be borne in mind, and that closing its gaps must depend upon chance preservations as well as on chance discoveries. A body of evidence bearing on the question of direct connection of Africa and South America, with much speculation derived from it, is supplied by the Permian reptiles of the order Mesosauria, and by certain members of the Triassic Rhynchocephalia, which are found in both continents. (See especially Edwin H. Colbert, 1952.) This finds support from the freshwater bivalves of the family Mutelidae, of which no fossils have been found in the northern hemisphere. Even in this case, the imperfections of the paleontological record and the possibility of convergence must be considered, for Myceiopoda dilucuU, described in 1921 by Pilsbry, from the Triassic of Pennsylvania, may belong to this family. The leadership among the group of Matthewsians has now somewhat naturally fallen to George Gaylord Simpson, who succeeded Matthew in the position of Curator in charge of Vertebrate Paleontology at the American Museum in 1944. Simpson had occasion to acquire the same kind of broad command of whole suc- cessions of extinct faunas. In a series of essays (1940-1952) he has dealt effec- tively with the difficulties of what appear as exceptional elements in these faunas — like the sudden appearance of the octodont rodents in South America, and the problems presented by the animal life of the West Indies and Madagascar. In 1945 he reviewed the whole classification of the mammals, living and extinct. Later Land-Bridge Speculations Matthew's reputation was so great among his close associates that it is dis- turbing to us to find him little recognized, or to find him even unknown in wide circles of geologists and geographers dealing with the problems of continental connections. Thus, in the face of isostasy, Charles Schuchert (1932) maintains connections from Madagascar to India, from Brazil to West Africa, and from Europe to Greenland. In an essay in the same year, with the title "Isthmian Links," Bailey Willis reduces the connections regarded as probable by Schuchert to narrow and sometimes tortuously crooked isthmuses, following the course of submarine ridges. Schuchert and Willis, on the geological side, thus stand solidly against the ideas of continental drift. Among botanists, one may cite W. H. Camp for his 1947 paper "Distribution Patterns in Modern Plants and the Problems of Ancient Dispersals." He thinks specifically in terms of Mesozoic east-west continent in the southern hemisphere, and more particularly of the origin of whole groups of flowering plants in the southern hemisphere. A European writer, Otto Wittmann, reviewing the problem in Zoogeographica, comes out flatly for a drifting back and forth of the continents (1934-1935). The case of the Hipparion bridge, a hypothetical Miocene land connection from Florida to Spain, including the Antilles and North Africa, is an especially flagrant example of post-Matthewsian irresponsibility, for it not only fails to consider the contradicting evidence, but is itself based on conflicting and erroneous interpretations of the fossils involved. Proposed in 1919 by L. Joleaud as a solution of the existence of the horse-like Hipparion in Florida and Spain in Miocene times, it presently was cited to explain all kinds of mammalian faunal relations between the Old AVorld and the New. By 1924 Joleaud had accepted 782 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES the idea of continental drift as a necessary alternative to the idea of transoceanic bridges; but he then finds himself forced to elaborate the drift theory to that of an "accordion movement back and forth" of the continental areas. Equally oblivious of the problems introduced by land connection theories is the French entomologist Rene Jeannel, who believes in a mid-Tertiary connection of the Mediterranean region with the West Indies as necessary to account for the dis- tribution of certain beetles (1935-1937) and later (1941) becomes a supporter of the ideas of continental drift as indicated by evidence from the faunas of the subantarctic islands like Kerguelen and the Crozets, as of continental drift in general from the viewpoint of entomology. It is quite evident that there are still numerous believers in the former existence of continents where the great oceans now are ; of movement of the conti- nents to their present positions from an original single continent; and of back and forth movement of "accordion-type" continents. There is a strong opposing school of conservatives, who hold to the belief that the continental platforms, though obviously often flooded by epicontinental seas, have been stable throughout the geological ages in which life has existed on land. Much of this controversy is primarily geological, and only secondarily zoogeographic. My concern in this matter has been lest the geologists base arguments on those of zoogeographers and that these then complete an argument in a circle by triumphantly pointing to the fact that the geologists support them. If the geological theories involved were restricted to pre-Paleozoic or even to Paleozoic times, zoogeographers could have little to say regarding them. Ecology and Animal Distribution Historical animal geography becomes properly scientific only when there is adequate positive evidence from paleontology as to the history in question, and negative evidence is doubly to be discounted because the fossil record is in itself incomplete, while our exploration of the world for what fossils have been pre- served is far from finished. The major factors producing disjunct distributions are the events of geological history, which are nonrecurrent. These can be recon- structed in convincing terms only when there is a quite exceptional wealth of fossil evidence. I have shown how wide is the divergence of opinion among zoo- geographers in this field. When we turn to contemporary animal distributions and examine them against the background of the existing environment, we enter a sharply contrasting realm of animal geography. Ecology, summing up existing environmental rela- tions, supplies the guiding principles in our studies when we turn to the geog- raphy of existing forms, especially at the species and infra-species level, and equally when we attend to the minor biotic geographic subdivisions of a continent. A new hypothesis can be tested by further observation, or even by experiment, and the whole field is subject to steady and logical growth with the advance of knowledge. The problems directly involved become ecological instead of historical. We still must face the enormous complexity of the total environment, but at least we can explore it at first hand. The ecological factors in animal distribution were appreciated more than a century ago, for we find Schmarda's division of the earth into regions based in SCHMIDT: ANIMAL GEOGRAPHY 783 part on considerations that are clearly ecological. In the development of this aspect of biogeography on scientific principles, the botanists took an early lead, and the whole field, for botany, was summarized by A. F. W. Schimper in his important book Pflanzengeographie auf physiologischer Grundlnge in 1898 (Eng. ed., 1903; 3d German ed. 1935). Thus ecological plant geography was summarized in an authoritative way more than a quarter-century before the appearance of the work by Richard Hesse, Tiei-geograpMe auf oekologischer Grundlage, which appeared in 1924. This has now been translated by myself, and quite completely revised by W. C. Alice and myself, for its successive American editions, 1937 and 1951, and it is still by no means as comprehensive a treatment of the ecological aspects of zoogeography as is Schimper 's work for phytogeography. If we search for the roots of ecological animal geography in North America, it is at once evident that they lie in the development of a systematic zoology focused on the existence of subspecies. As long as species were being described from isolated specimens, there might be very little knowledge of their actual geo- graphic ranges; but at the next level of analysis, the very idea of partitioning the species into subspecies required a definition of the ranges of both; and from such knowledge the step was easy to further ecological analysis of the meaning of the geographic ranges, and of analysis of the factors limiting such range. A school of description of subspecies grew up in North America hand in hand with the ambitious project of C. Hart Merriam for a biological survey of the continent. This in turn was directed into a broadly zoogeographie aspect by Merriams' development of the life zone theory (1890, 1898), in which the correspondence of altitude zones of mountains with the transcontinental climatic zones was pointed out, with an elaborate explanation of their temperature limitation. Merriam 's theory does not bear critical examination, though it was maintained for more than thirty years, and the work of the great United States Biological Survey was set in the Life Zone framework. Fortunately, the question of the existence of life zones is quite independent of the problems of their explana- tion. Merriam thought that northward distribution is limited by the sum of the positive temperatures (defined as degrees above an assumed physiological zero of 6°C) during the entire season of growth and reproduction, whereas southward limits were set by the mean temperature of a brief period during the hottest part of the year. That this was an extreme oversimplification is now evident; it is necessary to consider maximum and minimum temperatures; average temperature of the coldest part of the year; length and temperature of the frost- less season; amount of rainfall; degrees of atmospheric humidity and wind move- ment; day length microclimates; the considerable variety of edaphic factors; topographic barriers; and especially the complex of influences introduced by the sum of the favorable and unfavorable biotic factors. A thoroughgoing critique of the theory has been supplied by the botanists Livingston and Shreve (1921) and the zoologists Dice (1923), Kendeigh (1932), Shelf ord (1932), Dauben- mire (1938), and Pitelka (1941), to which list many more names might be added. The idea of temperature summation, originated as an ecological technique by Reaumur in 1735, is by no means to be discarded as useless — it remains as a measure of available heat supply in the growing season, and as one among the many factors that an ecological biogeographer must examine. Even the summed temperatures finally listed by Merriam have been found to 784 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES have included an overlooked fundamental error. How could a theory founded on so inadequate and one-sided an explanation of the facts have been so long main- tained and served so well as a guide for the exploration of North America? It is simply that the facts of the distribution of plant and animal life in a pattern related to the transverse climatic zones on a continental scale and to the altitude zones in mountains are quite independent of attempted explanations. It can scarcely be too much emphasized that animal life is dependent on plant life in far more ways than a diagram of food relations of plants and animals indicates. Animals live in a plant matrix, whose importance is measured somewhat by the difference in the tropical forest of the order of many thousands to one, and is still very great in the most densely populated savanna. I find this set forth force- fully by Alexander von Humboldt in his Kosmos (1845), and as early as 1808 in Ansichien der Natur. "Aspects of Nature" is in fact the key to the discrimination of the biotic areas for which we are in search; they prove to be precisely those vegetational areas that are visibly distinguishable in the landscape. The pattern of life zones, which is so conspicuously transcontinental in the open tundra and coniferous forest of northern North America, becomes more and more obscure in the southern half of the continent, and the Carolinas and California are radically distinct in both vegetation and fauna. The governing factors toward the south become more clearly those of humidity and rainfall instead of temperature, and there is then the further increase in complexity of the historical factors. We may accordingly turn to a much more complicated partition of the continent into biotic provinces. Work in this direction is exemplified in papers by Shelford and Pitelka, and especially by the little book by Lee R. Dice, The Biotic Provinces of North America (1943). Various papers by C. C. Adams deal with problems of animal geography from an ecological standpoint, and with his assistant at the University of Michi- gan, Alexander G. Ruthven (subsequently director of the University Museum), he focused interest on ecology in museum field work. His paper on the dispersal of the biota of the southeastern United States (1903) well illustrates these interests. Through Ruthven he left a permanent stamp on the Museum of Zoology of the University of Michigan, and Ruthven 's lectures on zoogeography influenced a generation of students. Distributions of the past were of course quite as much determined by the total ecology as those of the present; but we can only discern those paleoeco- logical factors with difficulty and with much more critical attention to the modes of occurrence of fossils than has been thought necessary hitherto. This is, how- ever, a fully recognized direction of effort in geology, and we have in progress a cooperative Treatise on 3Ia7'ine PaleoecoJogy of monumental proportions. Paleo- climatology, more specifically of the land areas, has had long attention, and is summarized in special works such as Brooks' Climate Through the Ages (2d ed. 1949). Two papers may be cited as exemplars of paleoclimatology applied to zoo- geographic studies. The first of these is Alfred Nehring's classic Ueher Tundren und Steppen der Jetzt- und Vorzeit (1890). Nehring analyzes the fauna of the existing northern tundra and of the existing Asiatic steppes (the semi-arid grasslands), and then examines fossil finds of these tundra and steppe animals in Europe in relation to the advances and retreats of the continental glaciers of the SCHMIDT: ANIMAL GEOGRAPHY 785 Pleistocene. In certain places there is a succession of deposits in which the life of the tundra (lemmings, reindeer, arctic fox) is found to be replaced by animals characteristic of the open steppe (such as lion, hyena, hamster, and jerboa), and this in turn by the remains of the aurochs and the modern fauna. He interprets this as reflecting the succession of climates and of vegetations associated with the retreat of the ice. In one of my own papers, I have been able to demonstrate a close parallel to the European westward extension of the steppes of Central Asia, in the eastward range of a part of our North American Great Plains fauna in the so-called "Prairie Peninsula" between the Great Lakes and the Ohio (Schmidt, 1937). Paleoclimatology in relation to the Pleistocene glaciations rests on abundant and conclusive evidence, with the extraordinary advantage in recent years of more exact dating by Carbon-14 analysis. As applied by Brooks and others to ancient climates, with different configurations of the continents, it has the same extremely speculative nature as the paleogeography on which it depends. At the very root of the problem of geographic range of the species we come upon the problems connected with the range of the individual animal, and more especially of the individual mated pair of breeding aggregations. Illuminating observations have been made on the establishment of well-defined territories in relation to their nests and to their feeding ground by birds, and these are being extended by critical observations to other groups of animals of the most diverse type. AVe are fortunate to have a summary of this important field of study by Mrs. Margaret Morse Nice "The Role of Territory in Bird Life" (1941), which includes a sketch of the history of the idea. The phenomenon is made conspicuous among birds by the songs of the males, which seem to be effective in establishing their spaced territories. The phenomenon in other animals may be complicated by the degree of social organization; the subject is summarized in The Principles of Animal Ecology (Alice et al., 1949). Island Life The problem of accounting for the life on islands in the sea has been close to the heart of animal geography from the earliest beginnings of thought. It became a crucial problem to Darwin in relation to giving up the doctrine of special creation in favor of a long-continued natural process. He at once set about making experiments on the viability of seeds after immersion in sea water; and began to assemble the observations of accidental transport of small animals by large, and of animals in general by floating vegetation, which he reports in the Origin of Species. Dispersal and capacity for dispersal are, in fact, basic to the whole of animal and plant geography. It is the denial of any capacity for transoceanic dispersal that lies at the root of the whole bridge-building contro- versy, and in part at the root of the arguments for continental drift. So con- vinced are the "connectionists" that there can be no overseas transport of land animals that the Galapagos Islands and the Hawaiian Islands have been thought to be quite as necessarily linked to the nearest continent as Britain and Borneo. It is one of the principal accomplishments in animal geography in recent decades to make a renewed analysis of such island life with the conclusion that the islands are indeed oceanic, and that the very existence of their land fauna proves its 786 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES capacity for overseas dispersal. The most notable advance in our understanding of such distributions has come with the appreciation of the importance of aerial dispersal by winds. The fact, for example, that the Hawaiian spider fauna con- sists exclusively of those families that disperse by means of gossamer flights is essentially conclusive as to their wind-blown origin. The complete renewed analysis of the Hawaiian fauna by Elwood C. Zimmerman confirms the dishar- monic nature of that fauna already evident in the earlier Fauna Hawaiiensis of R. C. L. Perkins. In his introductory volume for Insects of Hawaii (1948) Zimmerman presents a masterly and concliisive review of the problem of origin of the fauna. He finds it to be undoubtedly the growth of ages of flotsam- jetsam overseas immigration combined with wind-blown insects and accidental arrivals of strayed and off-course land birds. The West Indies have been a classical meeting ground for speculations as to the origin of their fauna and as to possible land connections with Central and South America. The much more radical idea that they had been directly con- nected by land with the Mediterranean region, via North Africa, as in Joleaud's llipparioyi bridge, crops up repeatedly. This is adopted by Jeannel to account for the presence of certain carabid beetles in the West Indies, namely of species whose congeners are found in the Atlantic Islands and North Africa or in the Old World generally. P. J. Darlington, Jr. (1938) has brought this problem into focus as a problem of dispersal; pointing out that the beetles cited by Jeannel as indicating land connection are among the smallest members of the carabid beetle fauna of the West Indies, which enormously increases the probability of their aerial dispersal. When the prevailing winds are plotted, and hurricanes and their tracks considered, the probabilities of arrival of these isolated elements of the West Indian fauna by aerial dispersal become overwhelmingly great. Darlington's discussion of aerial dispersal of insects is a return to Darwinian thinking about the problem of dispersal in general. A notable contribution to the problem of the possibilties and probabilities of aerial dispersal is supplied by direct studies of the objects found in the air by airplanes. The paper by P. A. (Hick, "The Distribution of Insects, Spiders, and Mites in the Air," published in 1939, summarizes data on collections made by means of special traps placed on airplane wings. More than 30,000 specimens of eighteen orders of insects, plus the spiders and mites, were obtained at altitudes ranging from 200 to 15,000 feet, the highest altitude being represented by a single specimen of spider. These observations establish beyond doubt the possibility of dispersal of small creatures of all kinds through the air. The most recent summary of the data of this kind is by Gislen, in 1948. Life of Fresh Waters Aerial dispersal had long before been shown to be the explanation of the strikingly wide distributions of freshwater organisms. Bodies of fresh water are usually sharply isolated, and by analogy with isolation on land, their life might be expected to exhibit great corresponding differences from lake to lake and from river to river, which is not found to be the case. The English geologist Thomas Belt further amplified the simple explanation of the wide uniformity of freshwater life offered by Darwin. He was much impressed, in Nicaragua, as SCHMIDT: ANIMAL GEOGRAPHY 787 Darwin had been in Brazil, by the radical difference in every aspect of land life from that familiar to him in England, and was the more astonished at the obvious similarities between the freshwater animals of tropical America and those of his native country. In The Naturalist in Nicaragua, published in 1874, he points out the fact that bodies of fresh water are in general relatively short-lived, at least in any geologic sense. This, on one hand, snuffs out the variations that develop, while on the other, it puts a premium on the capacity for dispersal. Many kinds of the smaller aquatic organisms have resting stages in which they may dry out and become a part of the dust blown up from a dried lake bottom or river bed. The great uniformity of the smaller animals of the fresh waters of North America and Europe is noteworthy even in the North American Great Lakes, which are no older than the last great advance of the continental glaciers. The effects of really long continued isolation in older bodies of water are thus all the more impressive and instructive, with many remarkable side lights on species formation. Lake Baikal in eastern Siberia, the Caspian Sea, Lake Tan- ganyika, and a very few others are preglacial in age, some perhaps dating from the mid-Tertiary; in each of these, animal life has evolved under strict isolation into wonderful series of endemic forms. We are fortunate to have a review of this subject by John Langdon Brooks (1950). To a somewhat lesser extent, the great river systems, like the Mississippi, the Danube, and the Yangtze are also ancient and isolated fresh waters, with strikingly peculiar animal types confined to them. The remnants of former lakes and river systems in old continental arid regions preserve an especially interesting record of origin and of subsequent isolation of their faunas. The phenomena of speciation in the fishes of the desert basins of the western United States have been summarized by Hubbs and Miller (1948). Another important advance in the study of the problems of dispersal has been made by analyzing the capacity for adjustment to brackish and salt water by freshwater fishes. It is evident that this capacity is much greater than has been suspected; and it is evident also that the limitation of the distribution of freshwater fishes by salt water barriers is a phylogenetic phenomenon; Myers (1949) formulated these ideas by grouping freshwater fishes according to their capacity for dispersal. His groups are: Primary, strictly intolerant of salt water; Secondary, rather strictly confined to fresh water, but relatively salt tolerant, at least for short periods; Vicarious, presumably nondiadromous freshwater rep- resentatives of primarily marine groups; Complementary, freshwater forms of marine groups, often diadromous, which become dominant in fresh water only when the first three divisions are absent; Diadromous fishes that migrate from fresh to salt water and vice versa; and Sporadic, for fishes that live and breed indifferently in salt or fresh water, without a fixed pattern of migration. Isolation and Speciation An ecological and evolutionary field of study that has been emphasized since the time of Darwin relates to the first beginnings of the origin of species. Darwin apparently greatly underestimated the role of geographic isolation in the sepa- ration of an incipient species from its parent form or from related forms, and this was first adequately emphasized by Moriz Wagner, the traveler and collector, whose attention was called to changes at the species level from one area to another 788 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES when a discernible barrier intervened. Wagner's papers on the topic (1868-1886) were collected by his nephew and published in 1889 under the title Die Entstehung der Arten durch rdumliche Sonderung. The importance of this aspect of animal geography is underlined in recent work in genetics and its relation to systematic botany, zoology, and paleontology. I need mention only Julian Huxley's The New Systematics, 1940, Ernst Mayr's comprehensive Systematics and the Origin of Species, 1942, and Allee, et al., The Principles of Animal Ecology, 1949. The phenomena of speciation are of especial interest in older bodies of fresh water and on older oceanic islands as may be seen in Brooks' and Zimmerman's papers cited above. Animal Geography of the Sea The vast and distinct field represented by the animal geography of the sea is in many respects extremely different from that of the land. Partition into faunal regions and provinces goes back to Schmarda (1853) and was thoughtfully re- viewed by Ortmann in 1896. Barriers are much less likely to be physiographic, are more likely to be related directly to temperature, and when physiographic, may also depend on mere distance. In one important respect, marine zoogeography is complementary to terrestrial zoogeography, namely in the analysis of the phenomenon of interruption of a land connection by the sea, when the new sea passage becomes a highway for dispersal of marine forms as soon as the land highway is broken. The separation of the continents of North and South America during the Tertiary involves a union in this area of the Atlantic and Pacific. The faunal relations thus produced were long ago pointed out by Jordan (1908); they are adequately summarized by Sven Elrnian in his Tiergeographie des Meeres (1935), of which a revised edition in English has recently appeared. Ekman sketches a most convincing picture of the major features of the world pattern of distribution of marine forms, of the operation of the open eastern Pacific and of the Atlantic as barriers to the coastal faunas, and of the historic importance of the Tethys Sea. Conclusion Animal geography is essentially an evolutionary study. It is only with diffi- culty definable as a separate science. In its descriptive branch it is one of the aspects of general natural history. If the contents of the eleven volumes on verte- brates of Brehm's Tierlehen were rearranged by geographic areas, we should have a comprehensive descriptive animal geography in eleven volumes. Interpretive zoogeography is so intimately related to ecology that it must always be considered as a branch of that synthetic science. Historical zoogeography, finally, is directly dependent on paleontology and thus in turn on geology. These intimate relations with several distinct sciences form the great merit of and the abundant justifica- tion for a science of animal geography. We have now long realized that speciali- zation must be balanced by synthetic sciences that bridge the gaps between the specialties and break down the barriers betw^een the scientists. It is good to turn from small to large problems, to take the long view, to think sometimes in terms of the world as a whole. Only then can we appreciate and interpret our own SCHMIDT: ANIMAL GEOGRAPHY 789 geography. And only thus may we attain the unstated but implicit goal of bio- logical studies — an understanding of "Man's Place in Nature." BIBLIOGRAPHY Adams, Charles C. 1903. Southeastern United States as a center of geographical distribution of iiora and fauna. Biol. Bull., 3:115-131. Airy, George Biddell 1856. An account of the pendulum experiments undertaken in the Harton Colliery for the purpose of determining the mean density of the earth. Philos. Trans. Roy. Soc. London, 1856:297-356. Allee, W. C, Alfred E. Emerson, Orlando Park, Thomas Park, and Karl P. Schmidt 1949. Principles of Animal Ecology, xii + 837 pp., 263 figs. Philadelphia: Saunders. Arldt, Theodor 1907. Die Entwicklung der Kontinente und ihrer Lebewelt, ein Beitrag zur ver- gleichenden Erdgeschichte. xxx + 729 pp., 17 text figs., 23 maps. (2nd ed., 1936, 1938.) Leipzig: Engelmann. 1919-1922. Handbuch der Palaeogeographie. 2 vols., illus. 1647 pp., 100 text figs. Leipzig: Borntraeger. Baker, H. B. 1912. The origin of continental forms, II. Ann. Rept. Mich. Acad. Sci., 1912:116-141. Belt, Thomas 1874. The Naturalist in Nicaragua: a Narrative of a Residence at the Gold Mines of Chontales; Journeys in the Savannahs and Forests with Observations on Animals and Plants in reference to the Theory of Evolution of Living Forms, xvi + 403 pp., illus. London: Murray. Brooks, C. E. P. 1949. Climate through the Ages: a Study of the Climatic Factors and Their Varia- tions. Rev. ed. 395 pp., 39 figs. New York: McGraw-Hill. Brooks, J. L. 1950. Speciation in ancient lakes. Quart. Rev. Biol., 25:30-60, 131-176, 8 figs. Camp, W. H. 1947. Distribution patterns in modern plants and the problems of ancient dispersals. Ecol. Monogr., 17:159-183, figs. 1-32. Chamberlin, T. C, and R. D. Salisbury 1904-1906. Geology. 3 vols., illus. New York: Holt. Chaney, Ralph W. 1947. Tertiary centers and migration routes. Ecol. Monogr., 17:139-148. Colbert, Edwin H. 1952. The Mesozoic Tetrapods of South America. Bull. Amer. Mus. Nat. Hist., 99: 237-254, figs. 27-28. Dana, James D. 1880. Manual of Geology: Treating of the Principles of the Science with Special Reference to American Geological History. 3rd ed. (1st ed. 1863). xiv + 911 pp., 12 pis., 1,159 text figs. (P. 797). New York: Ivison, Blakeman, Taylor. Darlington, P. J., Jr. 1938. The origin of the fauna of the greater Antilles, with discussion of dispersal of animals over water and through the air. Quart. Rev. Biol., 13:274-300, 5 figs. 790 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Daelington, p. J. Jr. (Cont.) 1948. The geographical distribution of cold-blooded vertebrates. Quart. Rev. Biol., 23:1-26, 105-123, 5 figs. Darwin, Charles R. 1859. Geographical distribution. Chapts. 12 and 13 in The Origin of Species, ix + 502 pp. London: Murray. Daubenmire, Rexford F. 1938. Merriam's life zones of North America. Quart. Rev. Biol., 13:327-332. Dice, L. R. 1923. Life zones and mammalian distribution. Journ. Mammal., 4:39-47. 1943. The biotic provinces of North America, viii + 78 pp., 1 map. Ann Arbor: Univ. Michigan Press. Du ToiT, Alexander L. 1937. Our Wandering Continents: an Hypothesis of Continental Drifting, xiii + 366 pp., 48 figs. London: Oliver and Boyd. Ekman, Sven 1935. Tiergeographie des Meeres. 542 pp., 244 figs. Leipzig: Akadem. Verlagsges. Forbes, Edward 1846. On the Connexion between the distribution of the existing Fauna and Flora of the British Isles, and the geological changes which have affected their area, especially during the epoch of the Northern Drift. Mem. Geol. Surv. Great Britain, 1:336-432. Forbes, H. 0. 1893. The Chatham Islands, their relation to a former southern continent. Roy. Geogr. Soc. Suppl. Pap. Ill, pp. 607-637. GiLMORE, C. W. 1943. Fossil lizards of Mongolia. Bull. Amer. Mus. Nat. Hist, 81:361-384, pis. 52, 22 text figs. GiSLEN, T. 1948. Aerial plankton and its condition of life. Biol. Rev., 23:109-126. Glick, p. a. 1939. The distribution of insects, spiders, and mites in the air. U. S. Dept. Agr. Tech. Bull., 673:1-151, 5 pis., 5 text figs. GtJNTHER, Albert 1858. On the geographical distribution of reptiles. Proc. Zool. Soc. London, 1858: 373-398. Handlirsch, Anton 1937. Neue Untersuchungen iiber die fossilen Insekten mit Erganzungen und Nach- tragen sowie Ausblicken auf phylogenetische, palaeogeographische und allge- mein biologische Probleme. Ann. Naturhist. Mus. Wien, 48:1-140. Haug, Emile 1922. Traits de geologic. 2 vols, (in 3), illus. Paris: Armand Colin. Hedley, Charles 1895. Considerations on the surviving refugees in austral lands of ancient antarctic life. Proc. Roy. Soc. New South Wales, 29:278-286. Heilprin, Angelo 1887. The Geographical and Geological Distribution of Animals, xii + 435 pp., 1 map. New York: Appleton. SCHMIDT: ANIMAL GEOGRAPHY 791 Hesse, Richard 1924. Tiergeographie auf oekologischer Grundlage. xii + 613 pp., 135 figs. Jena: Fischer. Hesse, Richard, W. C. Allee, and Karl P. Schmidt 1937. Ecological Animal Geography, xiv + 597 pp., 135 figs. New York: Wiley. Hooker, Joseph D. 1853. Flora Novae Zealandiae. Vol. II of The Botany of the Antarctic Voyage of H.M. Discovery-Ships "Erebus" and "Terror" in . . . 1839-1843 under the Command of Captain Sir J. C. Ross. 3 vols. London. Humboldt, Alexander von 1808. Ansichten der Natur, mit wissenschaftlichen Erlauterungen. viii + 344 pp. Tubingen. 1844. Kosmos. Entwurf einer physischen Weltbeschreibung. 4 vols. Stuttgart: Cotta. HUTTON, F. W. 1873. On the geographical relations of the New Zealand fauna. Trans. New Zealand Inst, 5:227-256. 1896. Theoretical explanations of the distribution of southern faunas. Proc. Linn. Soc. New South Wales, 21:36-47. Huxley, Julian (ed.) 1940. The New Systematics. viii + 583 pp., illus. Oxford: Oxford Univ. Press. Huxley, Thomas H. 1868. On the classification and distribution of the Alectoromorphae and Heteromor- phae. Proc. Zool. Soc. London, 1868:294-319, figs. 1-16, 1 map. 1870. Anniversary address of the president. Quart. Journ. Geol. Soc. London, 26: xxi-lxiv. Ihering, Hermann von 1891. On the ancient relations between New Zealand and South America. Trans. New Zealand Inst., 24:441-445. (Transl. from Das Ausland, No. 18, 1891.) Jeannel, Rene 1935. L'Archatlantis et le peuplement de la region mediteraneene. Arch. Mus. Hist. Nat. Paris, (6)12:413-426, 6 figs. 1941. Au Seuil de I'Antarctique. . . . Editions du Museum, No. 5. 236 pp., 16 pis., 27 text figs. Paris. Joleaud, L. 1924. L'histoire biogeographique de I'Amerique et la theorie de Wegener. Journ. Soc. Am6r. Paris, n.s., 16:325-360. Jordan, D. S. 1908. The law of geminate species. Amer. Nat., 42:73-80. Kendeigh, S. Charles 1932. A study of Merriam's temperature laws. Wilson Bull., 44:129-143, figs. 27-28. Marcou, Jules 1860. "Geographie antediluvienne." [Not seen.] In Raymond Furon, Le Jura dans les deux hemispheres. Carte globe a 1' epoque jurassique monstrant la dis- tribution des terres et des mers. Pp. 168-169. 1941. Matthew, W. D. 1915. Climate and Evolution. Ann. N. Y. Acad. Sci., 24:171-318, 33 figs. Reprinted as Spec. Publ. New York Acad. Sci., l:xi + 223 pp., 33 figs., 1939. 792 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Mavr, Ernst 1942. Systematics and the Origin of Species, xiv + 334 pp., 29 figs. New York: Colum- bia Univ. Press. Merriam, C. Hart 1890. Results of a biological survey of the San Francisco Mountain region and the desert of the Little Colorado, Arizona. North American Fauna, 3:1-136, pis. 1-13, maps 1-5. 1898. Life zones and crop zones of the United States. Bull. U. S. Dept. Agric, 10:7-79. Myers, George S. 1938. Fresh-water fishes and West Indian zoogeography. Ann. Rept. Smithson. Inst., 1937:339-364. 1949. Fresh-water fishes and East Indian zoogeography. Stanford Ichth. Bull., 4:11-21. 1949a. Salt-tolerance of fresh-water fish groups in relation to zoogeographical prob- lems. Bijdragen tot de Dierkunde, 28:315-322. Nehring, Alfred 1890. Ueber Tundren und Steppen der Jetzt- und Vorzeit, mit besonderer Beriick- sichtigung ihrer Fauna, viii -f 257 pp., illus. Berlin: Ferd. Diimmlers. Neumayr, Melchior 1886-1887. Erdgeschichte ... 2 vols., illus. Leipzig. Nice, Margaret Morse 1941. The role of territory in bird life. Amer. Midi. Nat, 26:441-487. Ortmann, a. E. 1896. Grundziige der marinen Tiergeographie. Anleitung zur Untersuchung der geo- graphischen Verbreitung mariner Tiere, mit besonderer Beriicksichtigung der Dekapodenkrebse. iv + 96 pp., 1 map. Jena: Fischer. 1901. The theories of the origin of the Antarctic faunas and floras. Amer. Nat., 35:139-142. OSUORN, H. F. 1900. The geological and faunal relations of Eui'ope and America during the Tertiary Period and the theory of successive invasions of an African fauna. Science, U.S., 11:561-574, 4 charts. Perkins, R. C. L. 1913. Introduction, being a review of the land-fauna of Hawaii. Fauna Hawaiiensis, l(pt. 6) :xv-ccviii, 16 pis. Pfeffer, Georg 1905. Die zoogeographischen Beziehungen Siidamerikas betrachtet an der Klassen der Reptilien, Amphibian, und Fische. Zool. Jahrb. Suppl., 8:407-442. Pitelka, Frank A. 1941, Distribution of birds in relation to major biotic communities. Amer. Midi. Nat., 25:113-137, figs. 1-11. Pratt, J. H. 1855. On the attraction of the Himalaya Mountains and of the elevated regions be- yond them upon the plumb line in India. Philos. Trans. Roy. Soc. London, 1855:53-100. Reaumur, Rene A. F. de 1735. Observations du thermometre. Alem. Acad. Roy. Sci., Paris, 1735:545-576. SCHMIDT: ANIMAL GEOGRAPHY 793 RUTIMEYER, C. I. 1867. Ueber die Herkunft unserer Tliierwelt. Eine zoogeographische Skizze. . . . Mit einem Verzeicliniss der fossilen und lebenden Scliweizerischen Saugethiere und einer Karte zur Andeutung der Geschichte der Tierverbreitung iru Allge- meinen. 57 pp., 1 map. Basel. SCHAKFF, R. F. 1909. On the evolution of continents as illustrated by the geographical distribution of existing animals. Proc. 7th Internat. Zool. Congr., pp. 1-14. Boston. SCHIMPER, A. F. W. 1898. Pflanzengeographie auf physiologischer Grundlage. xviii + 876 pp., 69 pis., 4 maps. Jena: Fischer. SCHMARDA, LUDWIG K. 1853. Die geographische Verbreitung der Thiere. viii + 755 pp., 1 map. Wien: C. Ceroid and Son. Schmidt, K. P. 1937. Herpetological evidence for the postglacial eastward extension of the steppe in North America. Ecology, 19:396-407, figs. 1-9. 1943. Corollary and commentary for "Climate and Evolution." Amer. Midi. Nat., 30:241-253. SCLATER, P. L. 1858. On "the general geographical distribution of the members of the class Aves. Journ. Linn. Soc, 2 (zool.) : 149-170. ScBiVENOR, J. B., I. H. BuRKiLL, Malcolm A. SMITH, A. Steven Corbet, H. K. Airy Shaw, P. W. Richards, and F. E. Zeuner. 1943. A discussion of the biogeographic division of the Indo-Australian Archipelago with criticism of the Wallace and Weber lines and of any other dividing lines and with an attempt to obtain uniformity in the names used for the divisions. Proc. Linn. Soc. London, 154 Sess., pp. 120-165, 2 figs. Shelford, Victor E. 1932. Life zones, modern ecology, and the failure of temperature summing. Wilson Bull., 44:144-157, figs. 29-34. Shuchert, Charles 1932. Gondwana Land Bridges. Bull. Geol. Soc. Amer., 43:875-916, pis. 24, 2 text figs. Simpson, George Gaylord 1940. Mammals and land bridges. Journ. Wash. Acad. Sci., 30:137-163, 6 figs. 1943. Turtles and the origin of the fauna of Latin America. Amer. Journ. Sci., 241: 413-429. 1943. Mammals and the nature of continents. Amer. Journ. Sci., 241:1-31. 1947. Holarctic mammalian faunas -and continental relationships during the Cenozoic. Bull. Geol. Soc. Amer., 58:613-687, figs. 1-6. 1950. History of the fauna of Latin America. Amer. Sci., 38:361-389, figs., 1-10. Suess, Eduard 1904-1924. The Face of the Earth. (Das Antlitz der Erde.) 5 vols., illus. (1st ed. 1883-1909). Oxford: Clarendon Press. Taylor. P. B. 1910. Bearing of the tertiary mountain belt on the origin of the earth's plan. Bull. Geol. Soc. Amer., 21:179-226. 794 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES Wagner, Moeiz 1889. Die Entstehung der Arten durch raumliche Sonderung. Gesammelte Aufsatze. 667 pp. Basel: Benno Schwabe. Wallace, Alfred Russell 1876. The Geographical Distribution of Animals with a Study of the Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earth's Surface. 2 vols., maps, illus. London: Macmillan. 1880: Island Life: or, the Phenomena and Causes of Insular Faunas and Floras, in- cluding a Revision and Attempted Solution of the Problem of Geological Climates, xvii + 526 pp., 3 maps, illus. London: Macmillan. Wegener, Alfred 1924. The Origin of Continents and Oceans, xx + 212 pp., 44 figs. London: Methuen. Willis, Bailey 1932. Isthmian links. Bull. Geol. Soc. Amer., 43:917-952, pis. 25-29. WiTTMANN, Otto 1934-1935. Die biogeographischen Beziehungen der Siidkontinente. Zoogeographica, 2:246-304, 12 figs.; 3:27-65, 10 figs. Zimmerman, Elwood C. 1948. Introduction. (Vol. I). In Insects of Hawaii, vii-xvii + 1-206 pp., 52 figs. Hono- lulu: Univ. Hawaii Press. THE CONSERVATION OF WILDLIFE By A. STARKER LEOPOLD Museum of Vertebrate Zoology, University of California, Berkeley In the year 1852 the Legislature of the new State of California passed a law protecting deer from hunting for six months of the year. This event marked the beginning of wildlife conservation in the West. In the year 1952 several branches of the California state government spent $3,379,000 on fish and game management (Calif. Dept. Fish and Game, 1953) and the state legislature had on its docket over 150 bills affecting wildlife in one way or another. During the intervening century there had evolved a com- plicated set of concepts and administrative programs concerned solely \vith ways and means of preserving and managing wild animal populations. Although the evolution of ideas in wildlife management is proceeding faster now than ever before, it may be well at this point to review the events of the past century as a guide to our future thinking and planning. Early Beginnings On the frontier, of course, there w^as little thought of wildlife preservation. Many a post mortem has been written about the careless treatment of animal resources by our pioneer forefathers. The slaughter of the bison and passenger pigeon, the ruthless commercialization of fur animals, the feather trade, have all been thoroughly lamented and there is no point in retracing the dark and bloody history here. In point of fact, human behavior is so completely condi- tioned by circumstances as to suggest that our most ardent conservationists today (the author included), had they been born into frontier society, would perhaps have acted much like their contemporaries. The concept of saving some- thing only assumes meaning when that thing becomes scarce. The conservation idea could not have been born until the native wealth of wildlife was clearly being dissipated. Our history of the conservation movement begins, therefore, with the first flickerings of recognition for its need. From 1852, when the first game-protective law was passed in California, until after 1900, despair over the steady shrinkage of game and fur resources in the West deepened into the conviction that game was ultimately doomed. More and more protective laws were dutifully adopted by the various state legislatures but there is no evidence of real optimism that the laws would stem the receding tide. Since no provision was made for enforcing the game laws, they were largely ignored by the hunting public, a fact well known to the legislators. The passage of game laws was in that era no more than an expres- sion of pious regret that the deer, elk, beavers, and so forth, were yielding to the inevitable advance of settlement. At the most it was hoped that legal pro- [795] 796 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES tection might serve to ration out the remaining stocks of game — to make the supply last a little longer. Then at the turn of the century there emerged a President with a firm con- viction that permanent preservation of wildlife and of sport hunting could be achieved as part of a general program of resource management. Theodore Roosevelt catalyzed an astonishing advance in conservation thinking. His long association with Gifford Pinchot and other foresters convinced him that wild crops such as forests and game could yield an annual harvest indefinitely if the rate of harvest were properly regulated and a basic breeding stock (or growing stock) were retained. Installed in the presidency in 1901, he deter- mined to make conservation his personal crusade. In the first years of his term, Roosevelt brought public attention to focus on the need for a broad national conservation program and he fanned to life the hope that such a program actu- ally could save some of the native beauty of the countryside. Scientists who had been busy cataloguing and describing the native animals suddenly came forth with a rash of pamphlets and magazine articles on preserving our vanish- ing wildlife. Newspapers devoted editorials and front page space to this new crusade. State legislatures from coast to coast busied themselves creating new conservation bureaus and departments. As a culmination, in 1908 Roosevelt invited all the Governors of the United States to a White House conference on conservation, and that event gave added stimulus to the movement which al- ready was well under way. The initial idea of the conservation movement was to protect and preserve the remnants of wildlife that had not been dissipated by the frontiersmen. Administrative programs were developed to implement various phases of the protective movement, and for the ensuing thirty years wildlife conservation implied in large part simple protection. The concept of managing and produc- ing game crops formulated slowly during this period. The three principal aspects of protection were (1) legal protection and law enforcement, (2) establishment of refuges and wildlife preserves, and (3) con- trol of natural predators. Legal Protection As stated above, game protective laws had little significance until wardens were sent into the field to enforce them. Most state w^arden forces were organized in the Rooseveltian era, at which time the hunting license was widely adopted as a device for financing law enforcement. Whereas many game-protective laws had been passed in the period from 1677, when Connecticut enacted the first such regulation, until 1852, when the custom reached California, and thousands more were adopted in the late nineteenth century, it was actually in the decade 1900- 1910 that effective legal protection was achieved. By then the near-extermination of some of the most numerous of native game species pointed up the great need for hunting control. Enthusiasm for game protection was stirred by the elo- quent writings of such leaders as William T. Hornaday (1913, 1914), William Butcher, Theodore S. Palmer, T. Gilbert Pearson, and John Phillips, and there quickly developed a strong public sympathy for the cause of protection, which of course was essential to its success. Almost from the start the protection movement brought demonstrable re- LEOPOLD: THE CONSERVATION OF WILDLIFE 797 suits in the way of game increases. Certain species such as deer and elk, which had been dangerously reduced, responded magnificently when the kill was limited, and this encouraged the conservationists in the firm conviction that they were on the right track. The fact that species whose habitats had been largely destroyed by agriculture and grazing did not respond equally (i.e., bison, prong- horn antelope, bighorn, waterfowl), was generally interpreted as a sign that protection was still inadequate. Efforts Avere redoubled to supply additional safeguards to these diminishing stocks. ^ Over the years the structure of legal game protection became more complex. Relatively simple laws were locally modified and refined, and license fees were raised to support constantly expanding warden forces. As an example of this evolution, table I traces the changes in seasons, bag limits, and special limita- tions on the hunting of deer in California. Although the table greatly simplifies legal details, it serves to show the growth of the protection program for a given species in one state. Table 1. A Chronological Summary of California Deer Hunting Regulations From 1852 to 1950 (from Longhurst, et ah, 1952) Bag limit Hunting Deer Year General seasons (Bucks) license tags Remarks 1852-82 — — — — Deer protected 6 months of year 1883-92 — — — — Antlerless deer protected 1893-94 Sept. 1-Oct. 15 — — — 1895-1900.. ..July 15-Oct. 15 — — — 1901-02 Aug. 1-Sept. 30 3 — — Night hunting and sale of meat prohibited 1903-04 July 15-Oct. 31 1905-06 Aug. 1-Oct. 15 1907-10 July 15-Sept. 30 1911-14 July 1-Oct. 31 1915-18 Aug. 1-Oct. 14 1919-20 Aug. 1-Oct. 14 1921-24 Aug. 1-Oct. 15 2, most of 1927-45 Aug. 1-Oct. 15 state; 1 2.00 $1.00 in part 2, most of 1946 Aug. 7-Oct. 21 state; 1 2.00 1.00 in part 2, most of 1947 Aug. 7-Oct. 15 state; 1 2.00 1.00 22 game districts in part 1, most of 1948 Aug. 7-Oct. 15 state; 2 3.00 1.00 in part 1, most of 1949-50 Aug. 7-Oct. 15 state; 2 3.00 1.00 Special seasons on antlerless deer in part in 3 localities 3 2 2 — — $1.00 — 2 1.00 — 6 game districts established 2 1.00 — 4 game districts 2 1.00 — 6 game districts; spike bucks protected 2 1.00 — Forked-horn bucks protected in northeastern district 798 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES By the late 1930 's and early 1940 's, game regulations were becoming so com- plex as to occupy an undue amount of time and attention on the agendas of state legislatures. In many states, plenary powers were transferred from gen- eral legislative bodies to fish and game commissions, which were much better able to cope with the details of regulating the game kill. Today most state laws dealing with wildlife are enacted by special commissions, but legislatures con- tinue to dabble in the field, which still has strong political significance. The ef- fort to remove wildlife administration and regulation from partisan politics has been only partly successful. Whereas the legal custodianship of most wildlife was vested in the individual states almost from the start of our national history, migratory birds became wards of the United States Government following ratification of a treaty with Canada in 1916 which established the international aspects of the problem. A similar treaty with Mexico was signed in 1937. Thus the federal government as- sumed an important responsibility in the legal protection of one class of game, and current regulations for the hunting of waterfowl and such other migra- tory birds as are still taken legally are set each year by the Secretary of Interior upon recommendation of the Director of the Fish and Wildlife Service. The complicated legal machinery, state and federal, that has been created to protect wildlife from overshooting has come to be considered in the public mind as the skeleton and backbone of wildlife conservation. Though it will al- ways be a necessary part of management, protection has probably been over- rated in importance, and now more recognition is being given to habitat man- agement as the principal key to game abundance. This changing point of view will be discussed in a later section. Refuges and Sanctuaries At the same time that general protection was extended to American wildlife in the form of legal restrictions on the kill, certain local areas were singled out for more intensive development as sanctuaries. The first of these was Lake Merritt in the City of Oakland, which was designated as a waterfowl sanctuary in 1870 by the California legislature. Subsequently nearly all of the states created numerous refuges embracing millions of acres, the assumption being that on these selected areas, game could flourish and spread out to surround- ing lands. On federal lands, wildlife was given complete protection on the national parks, although not in the first years of their existence. As early as 1864 the United States Congress set aside the Yosemite as a nature reserve but the main objective was to conserve the forest and scenery, not the wildlife. Finally, how- ever, in 1890 the Yosemite, General Grant, Sequoia, and Yellowstone areas, were formerly designated as national parks and in 1894 these areas were closed to hunting, as have been all the national parks and monuments ever since. Many state and municipal parks likewise are maintained as sanctuaries — a good policy in general, for animals that are both abundant and tame may be seen and enjoyed by visitors. Starting in 1903 the Federal government began withdrawing additional lands as wildlife refuges under the Bureau of Biological Survey, now the Fish LEOPOLD: THE CONSERVATION OF WILDLIFE 799 and Wildlife Service. Today there are 282 Federal wildlife refuges encompass- ing 18,500,000 acres (Day, 1949). One hundred and ninetj'-six of these areas are primarily maintained for migratory waterfowl; the balance, including some of the largest of the Federal refuges, serve to protect various species of upland game and colonial nesting birds. Supplementing the governmental refuges are many sanctuaries and pre- serves operated by municipalities, by conservation organizations such as the Na- tional Audubon Society, and by individual landowners. The refuge movement gained momentum during the early years of the pro- tective phase of game management and it reached a peak in the 1920's and 1930's. But as time went on it became clear that refuges per se were not the answer to the shortage of huntable game. For migratory waterfowl and for certain rare species of local occurrence the refuge is still and always will be a primary tool of management, but for upland game generally, closing some areas to hunting does not increase the level of game abundance in surrounding ter- rain. Most nonmigratory species are much too sedentary to "overflow" from a refuge and repopulate the rest of the countryside as had been postulated. Rather, the result of excluding hunters from parts of the game refuge serves merely to concentrate them in nonrefuge lands, thereby decreasing the avail- ability of game to the individual shooter. So the popularity of the refuge waned, and today most states are liquidating their refuge systems for upland game, though retaining those for waterfowl. Predator Control The third phase of the game protection program involved removal of preda- tory animals that were looked upon as "wicked citizens" of the wild community, destroying the breeding stocks that conservationists were striving to restore. Also, these same predators often preyed upon domestic livestock, rendering them doubly wicked in the public eye. And so the wolf and mountain lion, the coyote and bobcat, and many smaller offenders as well, came in for severe treatment. In addition to normal persecution by farmers, stockmen, and sportsmen, the predators were controlled systematically by special hunters, employed by the states and by the Federal government. Their demise was hastened in many localities by the payment of bounties or subsidies for scalps. Predator control proved generally to be the least satisfactory protective measure taken in behalf of game. Some of the large ungulates like deer and elk responded well enough to all this attention, but within fifty years they became so numerous in countless areas as to endanger their own forage supplies. Rigid hunting laws precluded effective control of populations by sportsmen, and removal of large predators such as wolves and lions had taken away the natural controls. In short, overenthusiasm for protecting game when it was scarce led ultimately to even more difficult problems that arose from an excess of game. The first great loss of deer by star- vation came on the Kaibab National Forest in Arizona, where a herd of a few thousand was built up to 100,000 in 1924 by rigid protection from hunting and predators. A plea by the Forest Service for reduction of the herd, to save the 800 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES range, was vigorously denied by state authorities, and in 1925 most of the deer died of starvation. Tlie herd continued to shrink until by 1939 there were only 10,000 left and the range was ruined. That sequence has been repeated a thousand times on deer ranges all over the West, the Lake States, and the North- east. Today many biologists argue in favor of less predator control on big game ranges as a safeguard against population irruptions. On the other hand, predator control as a measure to increase small game such as quail, pheasants, and rabbits, has never been proved to have any mate- rial effect at all. Great sums that have been paid for reduction of foxes, weasels, hawks, owls, and crows, probably have not raised the level of farm game above what the terrain would support anyway. Like many another sovereign remedy for game shortage, the control of preda- tors did not prove to be a panacea. Artificial Propagation — ^the Game Farm Mania After the program of wildlife protection was well under way, a new ap- proach was devised to give hunters more game to shoot. Various birds and mam- mals, some native but many exotic, were propagated in pens and liberated in the depleted coverts. The one great success of the restocking program was the introduction of the ring-necked pheasant from China into farmlands of the northern and central United States. Unfortunately, this initial coup de maitre inspired great confi- dence in propagation as a method of increasing game, leading over the years to expensive and usually fruitless attempts to repeat the process with other spe- cies. Most of the exotic pheasants, partridges, and grouse that were introduced failed to survive and the few that became established, such as the Chukar and Hungarian partridges, did so on a relatively small scale. Among native species, repeated studies have shown that pen-raised birds and mammals have a low survival rate and serve scarcely to augment the natu- ral crop of birds, raised in the wild at no expense. Where native stocks were literally exterminated by overhunting or trapping, introducing live-trapped animals from elsewhere often has been successful. For example, elk, antelope, and beaver have reoccupied great areas of range following reintroduction. But it was demonstrated that propagated stocks, for instance, of the wild turkey and the bobwhite quail were sometimes genetically inferior to native stocks in their ability to survive in the wild, and that mixing of the strains actually led to a decrease in wild populations. It has been proved that, where a breeding stock of game already exists, there is little advantage in attempting to build it up by artificial propagation. Habitat Management The shortcomings of simple protection and of propagation as methods of managing wildlife led finally to appreciation of the habitat as the transcend- ent force that, more than any other, determines the level of wild populations. It had long been recognized that each wild species was associated with a given sort of habitat and required certain types of food and of cover, but the idea LEOPOLD: THE CONSERVATION OF WILDLIFE 801 of producing game by the simple expedient of creating a suitable home for it was adopted slowly in this country. In the Old World, the purposeful preparation of the habitat for game had long since become standard practice. As early as the thirteenth century, Marco Polo noted that Kublai Khan maintained special preserves for partridges and pheasants, ''. . . for whose food the Great Khan caused millet, and other grains suitable to such birds, to be sown . . . every season, and gives strict command that no person shall dare to reap the seed; in order that the birds may not be in want of nourishment." The Khan likewise prepared special winter shelters and maintained a staff of gamekeepers to protect both the birds and their habi- tat. Marco Polo concludes: "In consequence of these attentions, he [the Khan] always finds abundant sport when he visits this country" [near Changanoor, Cathay] . At a somewhat later date in Europe, the planting of special coverts for pheasants and gray partridges became customary on country estates and on crown forests, and in Scotland the rotational burning of heather was found to be the least expensive and most effective way to increase numbers of red grouse. But these ideas were not carried to the New World. Much was said and written about preserving existing wildlife habitat, as for example on the na- tional forests, but cultural operations to create new or better habitat were not attempted until Herbert Stoddard, then with the United States Biological Sur- vey, undertook to study means of improving bobwhite shooting in Florida and Georgia. Stoddard's work on quail management in the Southeast was a mile- stone in American conservation. His book on The Bobwhite Quail (1931) sum- marized five years of intensive, scientific study of the bird in its natural environ- ment and pointed up the fact that the management of the land and its vege- tation had more to do with quail abundance than hunting, predators, or any other single factor. He showed how simple cultural operations could be used to create food and cover in proper interspersion, yielding a high density of quail and a high annual bag for hunters. Though the book is over twenty years old it is still the bible of game managers on Southern plantations. More im- portantly, it demonstrated the scientific approach to game production through good land management. At about the same time, another pioneer in the new era of wildlife manage- ment, Aldo Leopold, came forth with two volumes that reiterated Stoddard's findings and applied the basic tenets to game populations generally. The first of these, entitled Report on a Game Survey of the North Central States (1931) dealt with game conditions in one specific region. The second Game Management (1933), laid down the principles of scientific game production and harvest. From that time on, the study and the administration of wildlife resources was led gradually from the fields of politics and law into the fields of science and land management. The New Deal of the 1930's was a fruitful period in which scientific game management could grow. The federal government heavily subsidized conserva- tion projects of many kinds, and developing wildlife habitat became a recog- nized activity of such bureaus as the Soil Conservation Service, Forest Service, Tennessee Valley Authority, and Bureau of Land Management, as well as the Fish and Wildlife Service. State fish and game departments likewise began 802 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES spending money on game-range improvement instead of limiting their budgets to wardens, hatcheries, and game farms. The literature of the past decade has reflected this change in viewpoint. Among the important recent additions to the wildlife library are books by Gra- ham (1944, 1947), Trippensee (1948), Grange (1949), and Wing (1951), all of which emphasize the management and conservation of land and vegetation as the basis for game production. Perhaps of more fundamental importance than the administrative and lit- erary recognition of a scientific basis for game management has been the develop- ment of a body of trained professional men in the field. In 1937 a professional society was formed, called The "Wildlife Society, which began issuing a technical quarterly. The Journal of Wildlife Management. Many universities added trained wildlife men to their staffs, who in turn produced other trained men to fill administrative as well as academic positions. Wildlife management quickly assumed stature as a technical profession, comparable to forestry or the agri- cultural sciences. Wildlife Research Up to the time of Herbert Stoddard there was a decided separation between the scientific study of natural history and the administrative field of wildlife conservation. After Stoddard's demonstration of how scientific study could guide and orient conservation effort, wildlife research mushroomed into a thriv- ing field of activity. Some of the basic questions which have occupied wildlife students in the past and will continue to keep them busy for years into the future are : 1. Precisely what factors determine or limit wild populations? 2. How do various cultural operations of land, forestry, agriculture, graz- ing, etc.) affect game populations? 3. By what practicable means can game populations be increased? 4. What yields can and should be taken by hunters? These seemingly simple queries have proven to be very complex indeed. A digest of the considerable volume of data accumulated to date permits the fol- lowing tentative summary of the field of population dynamics which underlies the whole theory of management. Each wild population requires for its existence a number of indispensable components of habitat. These may be categorized under the headings (a) food, (b) cover, (e) water, and (d) special factors, such as grit, dusting facilities, salt, etc. If all of these are present in adequate amounts and in favorable juxta- position a population may exist. If the habitat is favorable, the density of the population will be higli. But if one or more of the environmental factors is limited in amount or in availability the population will tend to be less dense. The average level of a game population is therefore a function of the "carrying capacity" of the local habitat, a term used to express the sum effect of the en- vironment on the population. Thus, one area may support 50 deer per square mile but a similar tract with less forage might support only 30 deer per square mile. Food supply in the latter area is the factor limiting the population. Or, one quail population might be lower than another because cover or perhaps water was inadequate, hence limiting. LEOPOLD: THE CONSERVATION OF WILDLIFE 803 Whatever the level of a local game population, it reproduces annually and creates a "surplus," which in one way or another will be dispersed prior to the next breeding season. The surplus may be shot by hunters, or it may be taken by predators, or disease, or accidents. The important thing is that it will disap- pear. The annual increase in wild populations, which may be as low as 10 per cent in bears or as high as 300 per cent in quail, is never saved in a fully stocked habitat, but tends to be vulnerable to many kinds of losses, down to the level of "carrying capacity." Below that level, losses are few. These principles were first stated by Errington (1934, 1936, 1943) based on studies of bobwhites in Wisconsin and muskrats in Iowa. They have been verified by many subsequent investigations of other species in a variety of habitats and give support to the idea that management should strive to raise the carrying capacities of local environments as the cheapest and surest way to increase game. But these general concepts served merely to orient thinking without defining the specific nature of the relationships between vertebrate organisms and their environments. Recent research has sought to refine our understanding of "car- rying capacity" and of the reaction of individual animals to their surroundings and to each other. Considerable attention has been focused, for example, on the question of game nutrition. Even a few years ago, anything a bird or mammal ate was considered "food" and the only measure of importance applied to items of diet was quantitative. But it was noted that some types of food supported higher populations than others, and this led to investigations in qualitative nutrition. Among the various sorts of winter browse eaten by deer, for example, those species that seem best to maintain the animals have proved to be high in protein. On an adequate protein diet, deer remain strong and vigorous through the win- ter, the does bear many healthy fawns, and young adults breed at an early age. Conversely, on low protein the deer weaken, become subject to high losses from predators, disease, or outright starvation, and they raise few fawns (Longhurst et al., 1952). Thus food quality has a great deal to do with carrying capacity of deer ranges by regulating both the rate of increase and the extent of loss in the herds. Parallel studies of production and loss in quail populations suggest that there may be similar striking effects of changing quality in the diet. Another phase of ecology that is being much studied today is the matter of competition between members of a population and how competition serves to regulate population levels. Besides competing for food and for the best areas of cover, members of a dense population seem to affect each other in some subtle way that lowers reproductive rate. Thus in populations of bobwhite quail, ring- necked pheasant, mule deer, and brown rats, the rate of fecundity per individual female has been found to be inversely proportional to the density of the popu- lation. The most precise measure of this phenomenon has been made in rat populations in the city of Baltimore (Emlen et al., 1948; Davis, 1951). Following artificial reduction of the rats to a low level, there was a marked increase in the size and frequency of litters produced by the surviving females. As the population again approached carrying capacity, fecundity decreased until a stable population was restored, in which death rate and birth rate balanced . The obvious implication in management is that a heavy artificial kill, as by hunting, is compensated by an increased birth rate — a vital point in determining desirable rates of harvest. 804 A CENTURY OF PROGRESS IN THE NATURAL SCIENCES These examples serve to indicate the trends in current wildlife research. From this type of work on basic principles of population dynamics will inevit- ably come a better understanding of the critical or limiting factors that regu- late wild populations. Such knowledge in turn will guide future efforts in management. So rapid has been the progress in wildlife studies of the past decade that administrative procedures have been unable to keep pace with the changing ideas. Thus programs of predator control, artificial propagation, and close regu- lation of the kill that evolved over the past half-century are not easily aban- doned immediately upon discovery that there are better ways to expend available funds. Considerable investments in game farms and personnel trained in certain activities must be amortized and converted slowly to new undertakings. Like- wise, public opinion, which strongly influences legislation and administrative proceedings, must be reoriented periodically in line with scientific findings. Nevertheless, wildlife research in the past twenty years has had a tremendous influence on management policy, and that influence can be expected to grow in the future. "Wildlife and Land Use American land is being used more and more intensively to feed a nation that still is growing. Agriculture, grazing, forestry, and watershed protection are all primary uses of the land that in most areas will take precedence over wildlife production. If sport hunting is to be maintained as a form of outdoor recreation available to one and all, it will have to be carefully oriented to other forms of land use. Fortunately, game often may be produced in quantity on lands that are primarily dedicated to other uses. Thus, forest lands devoted to growing timber may, with only slight modification of management, also grow deer. Grain and pasture lands can produce a side crop of quail and pheasants. Meadows and sloughs can yield both beef and ducks. The task of wildlife research is to achieve an understanding of game populations and habitat relationships that will per- mit such dual planning of land use. The administrative task is to apply this knowledge. There are many practical difficulties to overcome in maintaining an optimum habitat for game on dual-use lands. Private landholders, for example, operate their farms and ranches primarily to produce marketable crops, and as yet there is no financial motive to spend time and money on habitat improvement for game. But many land practices that are of profit to the landowners also promote game crops. Fencing and planting gullies to prevent erosion creates coverts for wild- life as well. Many range practices that improve brushlands for cattle also benefit deer. Building farm ponds to conserve water for livestock and for irrigation creates habitat for ducks and some fur-bearers. Wildlife management is best sold to landowners then "via the back door" — as a secondary benefit of some profitable aspect of good farming. This places fish and game administrative bureaus in the position of being promoters of game production on lands not under their control. By subsidies and technical assistance they can induce a certain amount of habitat improve- ment on private lands. But the key man in the future of American wildlife will LEOPOLD: THE CONSERVATION OF WILDLIFE 805 continue to be the individual landowner. Recognition of this fact has led to increasing emphasis on conservation education and extension work among farm- ers and ranchers. The effective development of wildlife management on private lands is being seriously hindered by the legal machinery set up fifty years ago during the protective phase of game conservation. Ownership and custody of the game has been definitely placed by the courts in the hands of public agencies whose regulations governing hunting are in turn dictated in large part by organized sportsmen, not by the landowners who in fact are the real custodians of the game range. Rigid laws prevent the landowner from marketing a game crop in the way he markets his wheat or lambs, yet he is being asked to produce the crop for the public to harvest. Various legal devices such as cooperatives and licensed shooting preserves are now being tested to circumvent this problem, but with only partial success. Short seasons and unnecessarily conservative hunting laws still serve to discourage game management as a business enterprise on most private lands. In other words, there are traditional, educational, fiscal, and legal barriers to general application of research findings on how to produce game. On public lands the problem is relatively much simpler. For example, on the national forests game and fish production for public recreation is recognized as an important and in some areas as a primary use of the land. The Forest Service is not impelled solely by financial motives in establishing its land use policies, and where the public good is best served by devoting areas to wildlife (as for example, deer winter ranges, or reserves for rare species), conflicting uses may be excluded or made subservient. Noticeably more progress is being made in adopting scientific methods of game production on public lands than on private. It is clear, however, that on all lands throughout the nation there has been steady progress in adopting new and more effective methods of encouraging wildlife, and there is every reason to hope that substantial populations of shoot- able game, and of nongame native forms as well, can be retained despite intensi- fied use of land resources. Recognition of the importance of outdoor recreation in modern society has placed a premium on wildlife which will stimulate added effort among conservationists of the future. Summary Wildlife conservation in the United States started as an effort to preserve remnants of the native animal populations that had been severely depleted dur- ing the era of frontier exploitation. The initial stages were protective in nature and consisted principally of legal restrictions on hunting, setting aside refuges and sanctuaries, and controlling natural predators. After the protective program was well developed, a few trained biologists began to study game ecology in the field and learned that maintaining a suitable habitat for game was far more effective in sustaining wild populations than merely protecting existing breeding stocks. There followed a rapid reorienta- tion in conservation thinking and a parallel but slower adjustment in adminis- trative programs. One outgrowth of the success of the biological approach to wildlife manage- 806 ^ CENTURY OF PROGRESS IN THE NATURAL SCIENCES ment was the development of a technical profession, with training facilities in universities, accelerated research, and publication of scientific literature on game. In a very short time the nature of the profession changed from a quasi- legal and political undertaking to a scientific field comparable to forestry or the agricultural sciences. LITERATURE CITED California Department of Fish and Game 1953. Forty-second Biennial Report. 187 pp. Sacramento: Calif. State Printing Office. Davis, David E. 1951. A comparison of reproductive potential of two rat populations. Ecology, 32(3) : 469-475. Day, Albert M. 1949. North American Waterfowl, xx + 329 pp. New York: Stackpole and Heck, Inc. Emlen, J. T., Jr., A. W. Stokes, and C. P. Winsor 1948. The rate of recovery of decimated populations of brown rats in nature. Ecology, 29(2):133-145. Errington, Paul L. 1934. Vulnerability of bob-white populations to predation. Ecology, 15(2) :110-127. 1943. An analysis of mink predation upon muskrats in north-central United States. Iowa State Coll. Agri. Res. Bull. 320, pp. 797-924. Errington, Paul L., and F. N. Hamerstrom, Jr. 1936. The northern bob-white's winter territory. Iowa State Coll. Agri. Res. Bull. 201, pp. 301-443. Graham, Edward H. 1944. Natural Principles of Land Use. xiii + 274 pp. New York: Oxford Univ. Press. 1947. The Land and Wildlife, xiii + 232 pp. New York: Oxford Univ. Press. Grange, Wallace Bykon 1949. The Way to Game Abundance, with an Explanation of Game Cycles, xviil + 365 pp. New York: Charles Scribner's Sons. HoRNADAY, William T. 1913. Our Vanishing Wild Life: Its Extermination and Preservation, xvi + 411 pp. New York: Charles Scribner's Sons. 1914. Wildlife Conservation in Theory and Practice, vi + 240 pp. New Haven: Yale Univ. Press. Leopold, Aldo 1931. Report on a Game Survey of the North Central States. 299 pp. Madison, Wis.: Democrat Printing Co. 1933. Game Management, xxi -f 481 pp. New York: Charles Scribner's Sons. LoNGHURST, William M., A. Starker Leopold, and Raymond F. Dasmann 1952. A Survey of California Deer Herds, Their Ranges and Management Problems. Calif. Dept. Fish and Game, Game Bull. No. 6. 136 pp. Stoddard, Herbert L. 1931. The Bobwhite Quail: Its Habits, Preservation and Increase, xxix 4- 559 pp. New York: Charles Scribner's Sons. LEOPOLD: THE CONSERVATION OF WILDLIFE 807 Trippensee, Reuben Edwin 1948. Wildlife Management: Upland Game and General Principles, x + 479 pp. New York: McGraw-Hill Book Co., Inc. Wing, Leonard W. 1951. Practice of Wildlife Conservation, viii + 412 pp. New York: John Wiley and Sons, Inc.