XII A Century of Zoology in America
This article is intended as a brief survey of the development of zoology in America, and no attempt is made to give a general history of the science. There are numerous accounts in several languages of zoological history in general, among them being W. A. Locy’s “Biology and its Makers.” Brief outlines of the history of zoology may be found in many zoological and biological text-books.
For the history of American zoology the reader is referred to Packard’s report on “A Century’s Progress in American Zoology,” published in the American Naturalist, (10, 591, 1876), to Packard’s “History of Zoology,” published in volume 1 of the Standard Natural History (pp. lxii to lxxii, 1885); to G. B. Goode’s “Beginnings of Natural History in America,”172 and “Beginnings of American Science,”173 and to H. S. Pratt’s Manual of the Common Invertebrate Animals (pp. 1–9), 1916. In Binney’s “Terrestrial Air-breathing Mollusks of the United States” (1851) is a chapter on the rise of scientific zoology in the United States which well describes the zoological conditions in the early part of the century, while numerous monographs and papers give the history of the investigations on the various groups of animals or on special fields of study.
Brief biographical sketches of the most distinguished of our older Naturalists—Wilson, Audubon, Agassiz, Wyman, Gray, Dana, Baird, Marsh, Cope, Goode and Brooks are given in “Leading American Men of Science,” edited by David Starr Jordan, 1910. More extensive biographies have been published separately, and the activities of a number of the more prominent American zoologists have been recorded in the Biographical Memoirs of the National Academy of Sciences.
The developmental history of zoology in America falls naturally into four fairly well marked periods, namely:—1, Period of descriptive natural history, previous to 1847, embracing the early studies on the classification and habits of animals, characteristic of the zoological work previous to the arrival of Louis Agassiz in America. 2, Period of morphology and embryology, 1847–1870, during which the influence of Agassiz directed the zoological studies toward problems concerning the relationships of animals as indicated by their structure and developmental history. 3, Period of evolution, 1870–1890, when the principle of natural selection received general recognition and the zoological studies were largely devoted to the applications of the theory to all groups of animals. 4, Period of experimental biology, since 1890, during which time have occurred the remarkable advances in our knowledge of the nature of organisms through the application of experimental methods in the various branches of the modern science of biology.
American Zoology in 1818
At the beginning of the century which this volume commemorates, the accumulated biological knowledge of the world consisted mainly of what is to-day called descriptive natural history. The zoological treatises of the time were devoted to the names, distinguishing characters and habits of the species of animals and plants known to the naturalists of Europe either as native species or as the results of explorations in other parts of the world. This required little more than a superficial knowledge of their general anatomical structures.
The naturalists of those days had no conception of the life within the cell which we now know to form the basis of all the activities of animals and plants, nor had they even the necessary means of studying such life. The compound microscope, so necessary for the study of even the largest of the cells of the body, was not adapted to such use until 1835, although the instrument was invented in the seventeenth century. With the perfection of the microscope came a period of enthusiastic study of microscopic organisms and microscopic structures of higher animals and plants. It was not until twenty years after the founding of the Journal that the cell theory of structure and function in all organisms was established by the discoveries of Schleiden and Schwann.
The beginning of the nineteenth century saw great zoological activity in Europe, and particularly in France. Buffon’s great work on the Natural History of Animals had recently been completed, Cuvier had only one year before published his classic work in comparative anatomy, “Le Regne Animal,” and Lamarck’s “Philosophie Zoologique” had then aroused a new interest in classification and comparative anatomy from an evolutionary standpoint. E. Geoffroy St.-Hilaire was at the same time supporting an evolutionary theory based on embryonic influences resulting in sudden modifications of adult structure. These epoch-making discoveries and theories gained a considerable following in France, Germany and England, but seem to have had little influence on the zoological work of the following half century in America.
The science of zoology as understood to-day is commonly said to have been founded by Linnæus by the publication of the modern system of classification in the tenth edition of his “Systema Naturæ” in 1758. The influence of Linnæus aroused an interest in biological studies throughout Europe and stimulated new investigations in all groups of organisms. Such studies as related to animals naturally followed first the classification and relationship of species, that is, systematic zoology, and then led gradually into the development of the different branches of the subject, as morphology, comparative anatomy, physiology, and embryology, which eventually were recognized as almost independent sciences.
Of these sciences systematic zoology, which has come to mean the classification, structure, relationship, distribution and habits, or natural history, is the pioneer in any region. Thus we find in our new country at the time of the founding of the Journal in 1818, only sixty years after the publication of Linnæus’ great work, the beginning of American zoology taking the form of the collection and description of our native animals.
It is true that many of our more conspicuous and easily collected animals were described long before the opening of the nineteenth century, but this is to be credited mainly to the work of European naturalists who had made expeditions to this country for the purpose of studying and collecting. These collections were then taken to Europe and the results published there. We thus find in the 12th edition of Linnæus descriptions of over 500 American species, about half of which were birds. As an illustration of the extent to which some of these works covered the field even in those early days may be mentioned a monograph in two quarto volumes with many beautifully colored plates on the “Natural History of the rarer Lepidopterous Insects of Georgia.” This was published in London in 1797 by J. E. Smith from the notes and drawings of John Abbot, one of the keenest naturalists of any period.
During the early years of the nineteenth century, however, economic conditions in our country became such as to give opportunity for scientific thought. Educated men then formed themselves into societies for the discussion of scientific matters. This naturally led to the establishment of publications whereby the papers presented to the societies could be published and made available to the advancement of science generally. The most influential of these was the Journal of the Philadelphia Academy of Natural Science, which was established in 1817, and was devoted largely to zoological papers. The Annals of the New York Lyceum of Natural History date from 1823, and the Journal of the Boston Society of Natural History from 1834. The Transactions of the American Philosophical Society in Philadelphia and the Memoirs of the American Academy of Arts and Sciences in Boston also published many zoological articles.
In these publications and in the Journal, which was founded in 1818, appear the descriptions of newly discovered animal species, with observations on their habits.
The number of investigators in this field in the first quarter of the nineteenth century was but few, and most of these were compelled to take for the work such time as they could spare from their various occupations.
Gradually the workers became more numerous until about the middle of the century zoology was taught in all the larger colleges. The science thereby developed into a profession.
For some years the studies remained largely of a systematic nature, and embraced all groups of animals, but long before the close of the century the attention of the majority of the ever increasing group of zoologists was directed into more promising channels for research and there came the development of the sciences of comparative anatomy, physiology, embryology, experimental zoology, cytology, genetics, and the like, while the systematists became specialists in the various animal groups.
But the work in systematic zoology remains incomplete and many native species are still undescribed or imperfectly classified. It is perhaps fortunate that a few faithful systematists remain at their tasks and tend to keep the experimentalists from the disaster which might otherwise result from the confusion of the species under investigation.
Period of Descriptive Natural History — Previous to 1847
Of the few American naturalists whose writings were published toward the end of the eighteenth century and at the beginning of the nineteenth the names of William Bartram (1739–1823), Benjamin Barton (1766–1815), Samuel Mitchill (1764–1831), William Peck (1763–1822), and Thomas Jefferson (1743–1826), require special mention. Bartram’s entertaining volume describing his travels through the Carolinas, Georgia and Florida, published in 1793, contains a most interesting account of the birds and other animals which he found.
Barton wrote many charming essays on the natural history of animals, but was more particularly interested in botany. Mitchill’s most important works include a history of the fishes of New York (1814), and additions to an edition of Bewick’s General History of Quadrupeds. The latter, published in 1804, contains descriptions and figures of some American species and is the first American work on mammals.
Peck has the distinction of writing the first paper on systematic zoology published in America. This was a description of new species of fishes and was printed in 1794. He is also well known for his work on insects and fungi.
Jefferson in 1781 published an interesting book describing the natural history of Virginia, and during his presidency was of inestimable service to zoology through his support of scientific expeditions to the western portions of the country.
Previous to Agassiz’s introduction of laboratory methods of study in comparative anatomy and embryology in 1847, American naturalists generally confined their attention to the study of the classification and habits of the multitude of undescribed animals and plants of the region.
Such studies were naturally begun on the larger and more generally interesting animals such as the birds and mammals, and although many of these were fairly well described as to species before the opening of the nineteenth century, little was known of their habits. The natural history of our eastern birds first became well known through the accurate illustrations and exquisitely written descriptions of Alexander Wilson (in 1808–1813). Bonaparte’s continuation of Wilson’s work was published in four folio volumes beginning in 1826.
In 1828 appeared the first of Audubon’s magnificent folio illustrations of our birds. These were published in England, with later editions of smaller plates in America. Nuttall’s Manual of the Ornithology of the United States appeared in 1832–1834.
The second work on American mammals appeared in the second American edition of Guthrie’s Geography, published in 1815. The author is supposed to have been George Ord, although his name does not appear. In 1825 Harlan published his “Fauna Americana: Descriptions of the Mammiferous Animals inhabiting North America.” This was largely a compilation from European writers, particularly from Demarest’s Mammalogie, and had little value.
In 1826 Amos Eaton published a small “Zoological Text-book comprising Cuvier’s four grand divisions of Animals: also Shaw’s improved Linnean genera, arranged according to the classes and orders of Cuvier and Latreille. Short descriptions of some of the most common species are given for students’ exercises. Prepared for Rensselaer school and the popular class room.” “Four hundred and sixty-one genera are described in this text-book. They embrace every known species of the Animal Kingdom.” This is a compilation from European sources with a few American species of various groups included. On the other hand, Godman’s Natural History, in three volumes (1826–1828), was an illustrated and creditable work. Such was also the case with Sir John Richardson’s Fauna Boreali Americana of which the volume on quadrupeds was published in England in 1829. The other volumes on birds, fishes and insects appeared between 1827 and 1836. Audubon and Bachman’s beautifully illustrated “Quadrupeds of North America” was issued between 1841 and 1850.
About 1840 several of the states inaugurated natural history surveys and published catalogues of the local faunas. The reports on the animals of Massachusetts and New York are the most complete zoological monographs published in America up to that time. This is particularly true of DeKay’s Natural History of New York published between 1842 and 1844 in beautifully illustrated quarto volumes.
The leader in the systematic studies in the early part of the century was Thomas Say, who published descriptions of a large number of new species of animals, particularly reptiles, mollusks, crustacea and insects. Say’s conchology, printed in 1816 in Nicholson’s Cyclopedia, is the first American work of its kind. This was reprinted in 1819 under the title “Land and Fresh-water Shells of the United States.” In 1824–1828 appeared the three volumes of Say’s American Entomology.
The prominent position held by Say in the zoological work of this period is illustrated by the following paragraph from Eaton’s Zoological Text-book (1826, p. 133): “At present but a small proportion of American Animals, excepting those of large size, have been sought out... And though Mr. Say is doing much; without assistance, his life must be protracted to a very advanced period to afford him time to complete the work. But if every student will contribute his mite, by sending Mr. Say duplicates of all undescribed species, we shall probably be in possession of a system, very nearly complete, in a few years.” How different is the attitude of the zoologist of to-day who sees the goal much further away after a century’s progress through the industry of hundreds of investigators.
During the period of Say’s most active work he is reported to have “slept in the hall of the Philadelphia Academy of Natural Sciences, where he made his bed beneath the skeleton of a horse and fed himself on bread and milk.”
Next to Say, the most active zoologist of the early part of the century was Charles Alexander Lesueur, who described and beautifully illustrated many new species of fishes, reptiles, and marine invertebrates. A memoir by George Ord, published in this Journal (8, 189, 1849), gives a full list of Lesueur’s papers.
One of the most prolific writers of the period was Constantine Rafinesque, a man of great brilliancy but one whose imagination so often dominated his observations that many of his descriptions of plants and animals are wholly unreliable.
United States Exploring Expedition.—In 1838 a fortunate circumstance occurred which eventually brought American systematic zoology into the front ranks of the science. This opportunity was offered by the United States Exploring Expedition under the command of Admiral Wilkes. With James D. Dana as naturalist, the expedition visited Madeira, Cape Verde Islands, eastern and western coasts of South America, Polynesia, Samoa, Australia, New Zealand, Fiji, Hawaiian Islands, west coast of United States, Philippines, Singapore, Cape of Good Hope, etc.
Of the extensive collections made on this four-years’ cruise, Dana had devoted particular attention to the study of the corals and allied animals (Zoophytes) and to the crustacea. In 1846 the report on the Zoophytes was published in elegant folio form with colored plates. Six years later the first volume of the report on Crustacea appeared, with a second volume after two additional years (1854). These reports describe and beautifully illustrate hundreds of new species, and include the first comprehensive studies of the animals forming well-known corals. They remain as the most conspicuous monuments in American invertebrate zoology. Unfortunately the very limited edition makes them accessible in only a few large libraries. The other, equally magnificent, volumes include: Mollusca and Shells, by A. A. Gould, 1856; Herpetology, by Charles Girard, 1858; Mammalogy and Ornithology, by John Cassin, 1858.
Principal investigators.—Of the many writers on animals at this period of descriptive natural history, the following were prominent in their special fields of study:
Ayres, Lesueur, Mitchill, Storer, Linsley, Wyman, DeKay, Smith, Kirtland, Rafinesque and Haldeman described the fishes.
Green, Barton, Harlan, Le Conte, Say, and especially Holbrook, studied the reptiles and amphibia. Holbrook’s great monograph of the reptiles (North American Herpetology) was published between 1834 and 1845.
Wilson, Audubon, Nuttall, Cooper, DeKay, Brewer, Ord, Baird, Gould, Bachman, Linsley and Fox were among the numerous writers on birds.
Godman, Ord, Richardson, Audubon, Bachman, DeKay, Linsley and Harlan published accounts of mammals.
On the invertebrates an important general work entitled “Invertebrata of Massachusetts; Mollusca, Crustacea, Annelida and Radiata” was published by A. A. Gould in 1841, which contains all the New England species of these groups known to that date.
Lea, Totten, Adams, Barnes, Gould, Binney, Conrad, Hildreth, Haldeman, were the principal writers on mollusks. The crustacea were studied by Say, Gould, Haldeman, Dana; the insects by Say, Melsheimer, Peck, Harris, Kirby, Herrick; the spiders by Hentz; the worms by Lee; the coelenterates and echinoderms by Say, Mantell and others.
The history of entomology in the United States previous to 1846 is given by John G. Morris in the Journal (1, 17, 1846). In this article F. V. Melsheimer is stated to be the father of American Entomology, while Say was the most prolific writer. Say’s entomological papers, edited by J. L. Le Conte, were completely reprinted with their colored illustrations in 1859. The first economic treatise is that by Harris on Insects Injurious to Vegetation; printed in 1841. This has had many editions.
Zoology in the American Journal of Science, 1818–1846
The establishment of the Journal gave a further impetus to the scientific activities of Americans in furnishing a convenient means for publishing the results of their work. In the first volume of the Journal, for example, are two zoological articles by Say and a dozen short articles on various topics by Rafinesque, the latter being curious combinations of facts and fancy. Most of the zoological papers appearing in its first series of 50 volumes are characteristic of an undeveloped science in an undeveloped country. They deal, naturally, with observational studies on the structure and classification of species discovered in a virgin field, with notes on habits and life histories.
Many of the papers are purely systematic and include the first descriptions of numerous species of our mollusks, crustacea, insects, vertebrates and other groups. Of these, the writings of C. B. Adams, Barnes, A. A. Gould and Totten on mollusks, of J. D. Dana on corals and crustacea, of Harris on insects, of Harlan on reptiles, and of Jeffries Wyman and D. Humphreys Storer on fishes are representative and important.
The progress of zoology in America during the first twenty-eight years of the Journal’s existence, that is, up to the year 1846, is thus summarized by Professor Silliman in the preface to vol. 50 (page ix), 1847:
“Our zoology has been more fully investigated than our mineralogy and botany; but neither department is in danger of being exhausted. The interesting travels of Lewis and Clark have recently brought to our knowledge several plants and animals before unknown. Foreign naturalists are frequently visiting our territory; and, for the most part, convey to Europe the fruits of their researches, while but a small part of our own is examined and described by Americans: certainly this is little to our credit and still less to our advantage. Honorable exceptions to the truth of this remark are furnished by the exertions of some gentlemen in our principal cities, and in various other parts of the Union.”
During these 28 years the Journal had been of great service to zoology not only in the publication of the results of investigations but also in the review of important zoological publications in Europe as well as in America. There were also the reports of meetings of scientific societies. In fact all matters of zoological interest were brought to the attention of the Journal’s readers.
The Influence of Louis Agassiz
At the time of the founding of the Journal and for nearly thirty years thereafter descriptive natural history constituted practically the entire work of American zoologists. In this respect American science was far behind that in Europe and particularly in France. It was not until the fortunate circumstances which brought the Swiss naturalist, Louis Agassiz, to our country in 1846 that the modern conceptions of biological science were established in America.
Agassiz was then 39 years of age and had already absorbed the spirit of generalization in comparative anatomy which dominated the work of the great leaders in Europe, and particularly in Paris. The influence of Leuckart, Tiedemann, Braun, Cuvier and Von Humboldt directed Agassiz’s great ability to similar investigations, and he was rapidly coming into prominence in the study of modern and fossil fishes when the opportunity to continue his research in America was presented. On arriving on our shores the young zoologist was so inspired with the opportunities for his studies in the new country that he decided to remain.
Bringing with him the broad conceptions of his distinguished European masters, he naturally founded a similar school of zoology in America. It is from this beginning that the present science of zoology with its many branches has developed.
It must be remembered in this connection that the great service which Agassiz rendered to American zoology consisted mainly in making available to students in America the ideals and methods of European zoologists. This he was eminently fitted to do both because of his European training and because of his natural ability as an inspiring leader.
The times in America, moreover, were fully ripe for the advent of European culture. There were already in existence natural history societies in many of our cities and college communities. These societies not only held meetings for the discussion of biological topics, but established museums open to the public, and to which the public was invited to contribute both funds and specimens. This led to a wide popular interest in natural history. It was therefore comparatively easy for such a man as Agassiz to develop this favorable public attitude into genuine enthusiasm.
The American Journal of Science announces the expected visit of Agassiz as a most promising event for American Zoology (1, 451, 1846): “His devotion, ability, and zeal—his high and deserved reputation and... his amiable and conciliating character, will, without doubt, secure for him the cordial cooperation of our naturalists... nor do we entertain a doubt that we shall be liberally repaid by his able review and exploration of our country.” We of to-day can realize how abundantly this prophecy was fulfilled.
In the succeeding volume (2, 440, 1846) occurs the record of Agassiz’s arrival. “We learn with pleasure that he will spend several years among us, in order thoroughly to understand our natural history.”
Immediately on reaching Boston, Agassiz began the publication of articles on our fauna, and the following year he was appointed to a professorship at Harvard. The Journal says (4, 449, 1847): “Every scientific man in America will be rejoiced to hear so unexpected a piece of good news.” The next year the Journal (5, 139, 1848) records Agassiz’s lecture courses at New York and Charleston, his popularity with all classes of the people and the gift of a silver case containing $250 in half eagles from the students of the College of Physicians and Surgeons.
The service of Agassiz to American zoology, therefore, consisted not only in the publication of the results of his researches and his philosophical considerations therefrom, but also, and perhaps in even greater degree, in the popularization of science. In the latter direction were his inspiring lectures before popular audiences and the early publication of a zoological text-book. This book, published in 1848, was entitled “Principles of Zoology, touching the Structure, Development, Distribution and Natural arrangement of the races of Animals, living and extinct, with numerous illustrations.” It was written with the cooperation of Augustus A. Gould. The review of this book in the Journal (6, 151, 1848) indicates clearly the broad modern principles underlying the new era which was beginning for American zoology.
“A work emanating from so high a source as the Principles of Zoology, hardly requires commendation to give it currency. The public have become acquainted with the eminent abilities of Prof. Agassiz through his lectures, and are aware of his vast learning, wide reach of mind, and popular mode of illustrating scientific subjects... The volume is prepared for the student in zoological science; it is simple and elementary in style, full in its illustrations, comprehensive in its range, yet well considered and brought into the narrow compass requisite for the purpose intended.”
The titles of its chapters will show how little it differs in general subject matter from the most recent text-book in biology. Chapter I, The Sphere and fundamental principles of Zoology; II, General Properties of Organized Bodies; III, Organs and Functions of Animal Life; IV, Of Intelligence and Instinct; V, Of Motion (apparatus and modes); VI, Of Nutrition; VII, Of the Blood and Circulation; VIII, Of Respiration; IX, Of the Secretions; X, Embryology (Egg and its Development); XI, Peculiar Modes of Reproduction; XII, Metamorphoses of Animals; XIII, Geographical Distribution of Animals; XIV, Geological Succession of Animals, or their Distribution in Time.
A moment’s consideration of the fact that all these topics are excellently treated will show how great had been the progress of zoology in the first half of the nineteenth century. The sixty years that have elapsed since the publication of this book have served principally to develop these separate lines of biology into special fields of science without reorganization of the essential principles here recognized. This remained for many years the standard zoological and physiological text-book, and was republished in several editions here and in England. Another popular book is entitled “Methods of Study in Natural History” (1864).
More than 400 books and papers were written by Agassiz, over a third of which were published before he came to America. They cover both zoological and geological topics, including systematic papers on living and fossil groups of animals, but most important of all are his philosophical essays on the general principles of biology.
One of Agassiz’s greatest services to zoology was the publication of his “Bibliographia Zoologiæ et Geologiæ” by the Ray Society, beginning with 1848. The publication of the Lowell lectures in Comparative Embryology in 1849 gave wide audience to the general principles now recognized in the biogenetic law of ancestral reminiscence. As stated in the Journal (8, 157, 1849), the “object of the Lectures is to demonstrate that a natural method of classifying the animal kingdom may be attained by a comparison of the changes which are passed through by different animals in the course of their development from the egg to the perfect state; the change they undergo being considered as a scale to appreciate the relative position of the species.” These “principles of classification” are fully elucidated in a separate pamphlet, and are discussed at length in the Journal (11, 122, 1851).
One of the most interesting of Agassiz’s numerous philosophical essays, originally contributed to the Journal (9, 369, 1850), discusses the “Natural Relations between Animals and the elements in which they live.” Another philosophical paper contributed to the Journal discusses the “Primitive diversity and number of Animals in Geological times” (17, 309, 1854). Of his systematic papers, those on the fishes of the Tennessee river, describing many new species, were published in the Journal (17, 297, 353, 1854).
Agassiz’s beautifully illustrated “Contributions to the Natural History of the United States” cover many subjects in morphology and embryology, which are treated with such thoroughness and breadth of view as to give them a place among the zoological classics. The Essay on Classification, the North American Testudinata, the Embryology of the turtle, and the Acalephs are the special topics. These are summarized and discussed at length in the Journal (25, 126, 202, 321, 342, 1858; 30, 142, 1860; 31, 295, 1861).
The volume on the “Journey in Brazil” (1868) in joint authorship with Mrs. Agassiz is a fascinating narrative of exploration.
The conceptions which Agassiz held as to the most essential aim of zoological study are well illustrated in his autobiographical sketch, where he writes:174
“I did not then know how much more important it is to the naturalist to understand the structure of a few animals, than to command the whole field of scientific nomenclature. Since I have become a teacher, and have watched the progress of students, I have seen that they all begin in the same way; but how many have grown old in the pursuit, without ever rising to any higher conception of the study of nature, spending their life in the determination of species, and in extending scientific terminology!”
It is not surprising, then, that under such influence the older systematic studies should be replaced in large measure by those of a morphological and embryological nature.
The personal influence of Agassiz is still felt in the lives of even the younger zoologists of the present day. For the investigators of the present generation are for the most part indebted to one or another of Agassiz’s pupils for their guidance in zoological studies. These pupils include his son Alexander Agassiz, Allen, Brooks, Clarke, Fewkes, Goode, Hyatt, Jordan, Lyman, Morse, Packard, Scudder, Verrill, Wilder, and others—leaders in zoological work during the last third of the nineteenth century. Through such men as these the inspiration of Agassiz has been handed on in turn to their pupils and from them to the younger generation of zoologists.
The essential difference between the work of Agassiz and that of the American zoologists who preceded him was in his power of broad generalizations. To him the organism meant a living witness of some great natural law, in the interpretation of which zoology was engaged. The organism in its structure, in its development, in its habits furnished links in the chain of evidence which, when completed, would reveal the meaning of nature. Of all Agassiz’s pupils, probably William K. Brooks most fittingly perpetuated his master’s ideals.
Period of Morphology and Embryology, 1847–1870
The new aspect of zoology which came as a result of the influence of Agassiz characterized the zoological work of the fifties and sixties, that is, until the significance of the natural selection theory of Darwin and Wallace became generally appreciated.
The work in these years and well into the seventies was largely influenced by the morphological, embryological and systematic studies of Louis Agassiz and his school. The structure, development, and homologies of animals as indicating their relationship and position in the scheme of classification was prominent in the work of this period. The adaptations of animals to their environment and the application of the biogenetic law to the various groups of animals were also favorite subjects of study.
The most successful investigators in this period on the different groups of animals include:—Louis Agassiz on the natural history and embryology of coelenterates and turtles; A. Agassiz, embryology of echinoderms and worms; H. J. Clark, embryology of turtles and systematic papers on sponges and coelenterates; E. Desor, echinoderms and embryology of worms; C. Girard, embryology, worms, and reptiles; J. Leidy, protozoa, coelenterates, worms, anatomy of mollusks; W. O. Ayres and T. Lyman, natural history of echinoderms; McCrady, development of acalephs; W. Stimpson, marine invertebrates; A. E. Verrill, coelenterates, echinoderms, worms; A. Hyatt, evolutionary theories, bryozoa and mollusks; Pourtales, deep sea fauna; C. B. Adams, A. and W. G. Binney, Brooks, Carpenter, Conrad, Dall, Jay, Lea, S. Smith, Tryon, mollusks; E. S. Morse, brachiopods, mollusks; J. D. Dana, coelenterates and Crustacea; Kirtland, Loew, Edwards, Hagen, Melsheimer, Packard, Riley, Scudder, Walsh, insects; Gill, Holbrook, Storer, fishes; Cope, evolutionary theories, fishes and amphibia; Baird, reptiles and birds; J. A. Allen, amphibia, reptiles and birds; Brewer, Cassin, Coues, Lawrence, birds; Audubon, Bachman, Baird, Cope, Wilder, mammals.
The progress of ornithology in the United States previous to 1876 is well described in a paper by J. A. Allen in the American Naturalist (10, 536, 1876). A sketch of the early history of conchology is given by A. W. Tryon in the Journal (33, 13, 1862).
Jeffries Wyman was the most prominent comparative anatomist of this period. His work includes classic papers on the anatomy and embryology of fishes, amphibia, and reptiles.
Zoology in the American Journal of Science, 1846–1870
The fifty volumes of the second series of the Journal, including the years 1846 to 1870, cover approximately this period of morphology and embryology. During this period the Journal occupied a very important place in zoological circles, for J. D. Dana was for most of this period the editor-in-chief, while Louis Agassiz and Asa Gray were connected with it as associate editors. Moreover, in 1864 one of the most promising of Agassiz’s pupils, Addison E. Verrill, was called to Yale as professor of zoology and was made an associate editor in 1869.
In the Journal, therefore, may be found, in its original articles, together with its reports of meetings and addresses and its reviews of literature, a fairly complete account of the zoological activity of the period. The most important zoological researches, both in Europe and America, were reviewed in the bibliographic notices.
The most important series of zoological articles are by Dana himself. As his work on the zoophytes and crustacea of the U. S. Exploring Expedition continued, he published from time to time general summaries of his conclusions regarding the relationships of the various groups. Included among these papers are philosophical essays on general biological principles which must have had much influence on the biological studies of the time, and which form a basis for many of our present concepts.
The importance of these papers warrants the list being given in full. The titles are here in many cases abbreviated and the subjects consolidated.
General views on Classification, 1, 286, 1846.
Zoophytes, 2, 64, 187, 1846; 3, 1, 160, 337, 1847.
Genus Astraea, 9, 295, 1850.
Conspectus crustaceorum, 8, 276, 424, 1849; 9, 129, 1850; 11, 268, 1851.
Genera of Gammaracea, 8, 135, 1849; of Cyclopacea, 1, 225, 1846.
Markings of Carapax of Crabs, 11, 95, 1851.
Classification of Crustacea, 11, 223, 425; 12, 121, 238, 1851; 13, 119; 14, 297, 1852; 22, 14, 1856.
Geographical distribution of Crustacea, 18, 314, 1854; 19, 6; 20, 168, 349, 1855.
Alternation of Generations in Plants and Radiata, 10, 341, 1850.
Parthenogenesis, 24, 399, 1857.
On Species, 24, 305, 1857.
Classification of Mammals, 35, 65, 1863; 37, 157, 1864.
Cephalization, 22, 14, 1856; 36, 1, 321, 440, 1863; 37, 10, 157, 184, 1864; 41, 163, 1866; 12, 245, 1876.
Homologies of insectean and crustacean types, 36, 233, 1863; 47, 325, 1894.
Origin of life, 41, 389, 1866.
Relations of death to life in nature, 34, 316, 1862.
Of the above, the articles on cephalization as a fundamental principle in the development of the system of animal life have attracted much attention. The evidence from comparative anatomy, paleontology, and embryology alike supports the view that advance in the ontogenetic as well as in the phylogenetic stages is correlated with the unequal growth of the cephalic region as compared with the rest of the body. Dana shows that this principle holds good for all groups of animals. His homologies of the limbs of arthropods and vertebrates, however, do not accord with more modern views.
Other papers on the same and allied topics were published by Dana in other periodicals. His most conspicuous zoological works, however, are his reports on the Zoophytes and Crustacea of the United States Exploring Expedition, 1837–1842. The former consists of 741 quarto pages and 61 folio plates, describing over 200 new species, while the Crustacea report, in two volumes, has 1620 pages and 96 folio plates, with descriptions of about 500 new species. Each of these remains to-day as the most important contribution to the classification of the respective groups. The relationships of the species, genera and families were recognized with such remarkable judgment that Dana’s admirable system of classification has remained the basis for all subsequent work.
Dana’s critical reviews (25, 202, 321, 1858) of Agassiz’s “Contribution to the Natural History of the United States” are among the most interesting of his philosophical discussions concerning the relationships of animals as revealed by their structure, their embryology, and their geological history.
The remaining zoological articles in this series cover nearly the whole range of systematic zoology. Especially important are the articles by Verrill on coelenterates, echinoderms, worms and other invertebrates.
In the years following the publication of Darwin’s Origin of Species in 1859 occur many articles on the theory of natural selection. Some of the writers attack the theory, while others give it more or less enthusiastic support.
Experimental methods in solving biological problems were little used at this time, although a few articles of this nature appear in the Journal. Of these, a paper by W. C. Minor (35, 35, 1863) on natural and artificial fission in some annelids has considerable interest to-day.
Exploring Expeditions
Of the important zoological expeditions the following may be selected as showing their influence on American Zoology:
The North Pacific Expedition, with William Stimpson as zoologist, returned in 1856 with much new information concerning the marine life of the coasts of Alaska and Japan and many new species of invertebrates.
In 1867–1869 the United States Coast Survey extended its explorations to include the deep sea marine life off the southeastern coasts and Gulf of Mexico under the leadership of Pourtales and Agassiz.
The Challenger explorations (1872–1876) added greatly to the knowledge of marine life off the American coast as well as in other parts of the world.
The explorations of the United States Fish Commission succeeded those of the Coast Survey in the collection of marine life off our coasts and in our fresh waters. These have continued since 1872 and have yielded most important results from both the scientific and economic standpoints.
Under the charge of Alexander Agassiz the Coast Survey Steamer “Blake,” in 1877 to 1880, was engaged in dredging operations in three cruises to various parts of the Atlantic. The U. S. Fish Commission Steamer “Albatross,” also in charge of Agassiz, made three expeditions in the tropical and other parts of the Pacific in the years from 1891 to 1905. The study of these collections has added greatly to our knowledge of systematic zoology and geographical distribution. The reports on some of the groups are still in course of preparation.
Period of Evolution, 1870–1890
The time from 1870 to 1890 may be appropriately called the period of evolution, for although it commences eleven years after the publication of the Origin of Species, the importance of the natural selection theory was but slowly receiving general recognition. The hesitation in accepting this theory was due in no small degree to the opposition of Louis Agassiz. After the acceptance of evolution, although morphological and embryological studies continued as before, they were prosecuted with reference to their bearing on evolutionary problems.
Following closely the methods which had produced so much progress during the life of Agassiz, the field of zoology was now occupied by a new generation, among whom the pupils of Agassiz were the most prominent.
The teaching of biology at this time was also strongly influenced by Huxley, whose methods of conducting laboratory classes for elementary students were adopted in most of our large schools and colleges. This placed biology on the same plane with chemistry as a means for training in laboratory methods and discipline, with the added advantage that the subject of biology is much more intimately connected with the student’s everyday life and affairs.
This increasing demand for instruction in biology and the consequent necessity for more teachers brought an increasing number of investigators into this field.
Conspicuous in this period was the work of E. D. Cope, best known as a paleontologist, but whose work on the classification of the various groups of vertebrates stands preeminent, and whose philosophical essays on evolution had much influence on the evolutionary thought of the time. He was a staunch supporter of the Lamarckian doctrine. Alpheus Hyatt also maintained this theory, and brought together a great accumulation of facts in its support. He thereby contributed largely to our knowledge of comparative anatomy and embryology. A. S. Packard, whose publications cover a wide range of topics, was best known for his text-books of zoology and his manuals on insects.
W. K. Brooks was a leading morphologist and embryologist. S. F. Baird, for many years the head of the United States Fish Commission, was the foremost authority on fish and fisheries and is also noted for his work on reptiles, birds and mammals. The man of greatest influence, although by no means the greatest investigator, was C. O. Whitman. It is to him that we owe the inception of the Marine Biological Laboratory, the most potent influence in American zoology to-day; the organization of the American Morphological Society, the forerunner of the present American Society of Zoologists; and the establishment of the Journal of Morphology. G. B. Goode was distinguished for his work on fishes and for his writings on the history of science.
E. L. Mark, C. S. Minot, and Alexander Agassiz were acknowledged leaders in their special fields of research—Mark in invertebrate morphology and embryology, and Minot in vertebrate embryology, while Alexander Agassiz made many important discoveries in the systematic zoology and embryology of marine animals, and to him we owe in large measure our knowledge of the life in the oceans of nearly all parts of the world.
The knowledge of the representatives of the different divisions of the American fauna had now become sufficient to allow the publication of monographs on the various classes, orders and families. At this time also particular attention was given to the marine invertebrates of all groups.
Of the many investigators working on the various groups of animals at this time only a few may be mentioned. The protozoa were studied by Leidy, Clark, Ryder, Stokes; the sponges by Clark, Hyatt; the coelenterates by A. Agassiz, S. F. Clarke, Verrill; the echinoderms by A. Agassiz, Brooks, Kingsley, Fewkes, Lyman, Verrill; the various groups of worms by Benedict, Eisen, Silliman, Verrill, Webster, Whitman; the mollusks by A. and W. G. Binney, Tryon, Conrad, Dall, Sanderson Smith, Stearns, Verrill; the Brachiopods by Dall and Morse; the Bryozoa by Hyatt; the crustacea by S. I. Smith, Harger, Hagen, Packard, Kingsley, Faxon, Herrick; the insects by Packard, Horn, Scudder, C. H. Fernald, Williston, Norton, Walsh, Fitch, J. B. Smith, Comstock, Howard, Riley and many others; spiders by Emerton, Marx, McCook; tunicates by Packard and Verrill; fishes by Baird, Bean, Cope, Gilbert, Gill, Goode, Jordan, Putnam; amphibians and reptiles by Cope; birds by Baird, Brewer, Coues, Elliott, Henshaw, Allen, Merriam, Brewster, Ridgway; and the mammals by Allen, Baird, Cope, Coues, Elliott, Merriam, Wilder.
Interest in the evolutionary theory continued to increase and eventually developed into the morphological and embryological studies which reached their culmination between 1885 and 1890 under the guidance of Whitman, Mark, Minot, Brooks, Kingsley, E. B. Wilson and other famous zoologists of the time. In these years the Journal of Morphology was established and the American Morphological Society was formed.
The morphological, embryological and paleontological evidences of evolution as indicated by homologies, developmental stages and adaptations were the most absorbing subjects of zoological research and discussion.
Zoology in the American Journal of Science, 1870–1918
The third series of the Journal (1870–1895), likewise including fifty volumes, embraces this period of zoological activity in morphological and embryological studies, culminating with the inception of the modern experimental methods.
In this period also occurred the greatest progress in marine systematic zoology, due to the explorations of the United States Fish Commission off the Atlantic Coast. The Journal had an important share in the zoological development of this period also, for A. E. Verrill, who was now an associate editor, was in charge of the collections of marine invertebrates. Consequently most of the discoveries in this field were published in the Journal in numerous original contributions by Verrill and his associates. The explorations of the U. S. Fish Commission Steamer “Albatross” are described from year to year by Verrill, with descriptions of the new species of invertebrates discovered.
The numerous original contributions by Verrill on subjects of general zoological interest as well as on those of a systematic nature give this third series of the Journal much zoological importance. Verrill’s papers cover almost the whole field of descriptive zoology, but are mainly devoted to marine invertebrates. Those which were originally contributed to the Journal or summarized by him in his literature reviews include the following topics:
Sponges, 16, 406, 1878.
Coelenterates, 37, 450, 1864; 44, 125, 1867; 45, 411, 186; 46, 143, 1868; 47, 282, 1869; 48, 116, 419, 1869; 49, 370, 1870; 3, 187, 432, 1872; 6, 68, 1873; 21, 508, 1881; 6, 493, 1898; 7, 41, 143, 205, 375, 1899; 13, 75, 1902.
Echinoderms, 44, 125, 1867; 45, 417, 1868; 49, 93, 101, 1870; 2, 430, 1871; 11, 416, 1876; 49, 127, 199, 1895; 28, 59, 1909; 35, 477, 1913; 37, 483, 1914; 38, 107, 1914; 39, 684, 1915.
Worms, 50, 223, 1870; 3, 126, 1872.
Mollusks, 49, 217, 1870; 50, 405, 1870; 3, 209, 281, 1872; 5, 465, 1873; 7, 136, 158, 1874; 9, 123, 177, 1875; 10, 213, 1875; 12, 236, 1876; 14, 425, 1877; 19, 284, 1880; 20, 250, 251, 1880; 2, 74, 91, 1896; 3, 51, 79, 162, 355, 1897.
Crustacea, 44, 126, 1867; 48, 244, 430, 1869; 25, 119, 534, 1908.
Ascidians, 1, 54, 93, 211, 288, 443, 1871; 20, 251, 1880.
Dredging operations and marine fauna, 49, 129, 1870; 2, 357, 1871; 5, 1, 98, 1873; 6, 435, 1873; 7, 38, 131, 405, 409, 498, 608, 1874; 9, 411, 1875; 10, 36, 196, 1875; 16, 207, 371, 1878; 17, 239, 258, 309, 472, 1879; 18, 52, 468, 1879; 19, 137, 187; 20, 390, 1880; 22, 292, 1881; 23, 135, 216, 309, 406, 1882; 24, 360, 477, 1882; 28, 213, 378, 1884; 29, 149, 1885.
Miscellaneous, 39, 221, 1865; 41, 249, 268, 1866; 44, 126, 1867; 48, 92, 1869; 3, 386, 1872; 7, 134, 1847; 10, 364, 1875; 16, 323, 1878; 20, 251, 1880; 3, 132, 135, 1897; 9, 313, 1900; 12, 88, 1901; 13, 327, 1902; 14, 72, 1902; 15, 332, 1903; 24, 179, 1907; 29, 561, 1910.
S. I. Smith describes the metamorphosis of the Crustacea (3, 401, 1872; 6, 67, 1873), species of crustacea (3, 373, 1872; 7, 601, 1874; 9, 476, 1875), and dredging operations in Lake Superior (2, 373, 448, 1871). In this series occurs also a series of papers on comparative anatomy and embryology from the Chesapeake Zoological Laboratory in charge of W. K. Brooks. In the 39th and 40th volumes of the third series (1890) occur several papers on evolutionary topics by John T. Gulick (39, 21; 40, 1, 437) which have attracted much attention.
Before the end of this period, however, the Journal was relieved from the necessity of publishing zoological articles by the establishment of several periodicals devoted especially to the various fields of zoology. We find, therefore, but few exclusively zoological papers after 1885, although articles of a general biological interest and the reviews of zoological books continue.
In the fourth series of the Journal, beginning in 1896, occur also a number of articles on systematic zoology by Verrill and others and several papers having a general biological interest. Brief reviews of a small number of zoological books are still continued, but at the present day the Journal, which played so important a part in the early development of American zoology, has been given over to the geological and physical sciences in harmony with the modern demand for specialization.