Museum of Zoology

College of Literature, Science, and the Arts

The University of Michigan: An Encyclopedic Survey was first published beginning in 1942. For its 2017 Bicentennial, the University undertook the most significant updating of the Encyclopedia since the original, focusing on academic units. Entries from all versions are compiled in the Bicentennial digital and print-on-demand edition. Contents

1. The Museum of Zoology (1942) 1 J. Speed Rogers

2. The Museum of Zoology (2015) 30 Diarmaid Ó Foighil

[1]

The Museum of Zoology (1942)

J. Speed Rogers

THE Museum of Zoology is a research and teaching unit of the University for the study of the evolution, distribution, and systematic relationships of animals. Its collections comprise some 50,000 mammals, 110,000 birds, 130,000 reptiles and amphibians, 2,100,000 fish, 2,500,000 insects, and 2,000,000 mollusks, with the accompanying data essential for study. The collections are as notable for the wealth and accuracy of their field data as for the excellence of their representation of the respective groups, and they have given the Museum an international reputation as a center for graduate teaching and research in natural history. The Museum occupies the first, second, and third floors and much of the basement in the north wing of the University Museums Building. The collections are housed in fireproof ranges designed for safe storage and ready access to any individual specimen or desired series. Complementing the ranges are staff offices and study rooms, aquarium and live- rooms, preparation rooms, divisional libraries, and a seminar and classroom, an over-all area of some 46,000 square feet. 2 Museum of Zoology

The staff includes a director and eleven curators, as well as research and student assistants, an artist for the illustration of its publications, and a small secretarial staff. The curators and the director have academic status in the Department of Zoology, and their duties are more or less equally divided between curatorial work, research, and teaching. These functions, indeed, are not separable in any real sense. The extensive collections and the data that accompany them are the essential tools for both research and teaching, and so long as they are in use require a continuing curatorial process which, if it is at all competent, depends upon current as well as completed research. The teaching is largely but not exclusively graduate. In addition to directing the doctoral studies of some thirty to forty students, staff members give courses and seminars in mammalogy, ornithology, herpetology, ichthyology, entomology, malacology, and systematics and supervise individual studies, which any qualified student may elect. The Museum of Zoology is one of the University’s oldest units for specialized research. Although its official existence as a separate unit dates from as late as 1912-13, its modern existence began in 1903, as a direct descendant of the University Museum of Natural History provided for in the enabling act of 1837. The transformation of the old Cabinet and later Museum of Natural History into a Museum of Zoology came in large part from the cumulative enthusiasm and efforts of three great naturalists who were successively in charge of the zoological part of the natural history collections. The first, Professor Joseph Beal Steere (’68, ’70l, Ph.D. hon. ’75), Curator of the Museum from 1876 to 1894, was a pioneer student of animal geography. His zoological explorations in South America, the Philippines, and parts of the Dutch East Indies brought the first large accessions of study materials as contrasted with those for exhibit, and gave the Museum of Natural History a strong zoological bent. The second man, Charles Christopher Adams (Illinois Wesleyan ’95, Ph.D. Chicago ’06, Sc.D. Illinois Wesleyan ’20), was brought to Michigan in 1903 by Jacob Reighard, Steere’s successor as Professor of Zoology, to be Curator in charge of the Museum, The Museum of Zoology (1942) 3 because Reighard’s own interests were in other fields. Adams was one of America’s pioneer students of ecology and faunal relationships, and it was due to his interest that attention was turned to a detailed study of the Michigan fauna and an analysis of its ecological and geographic history and relationships. He instituted a series of faunistic and ecological surveys of the state and stressed the fact that abundant specimens and detailed field data are essential for the development of scientific natural history. The third man was Alexander Grant Ruthven (Morningside ’03, Ph.D. Michigan ’06, LL.D. California ’38, D.H.C. Catholic University of Chile ’44), Adams’ student and chief lieutenant in the zoological explorations of the state and in 1906 his successor as Curator of the University Museum. Ruthven’s own researches related the data and problems of geographic distribution with those of ecological adaptation. He conceived of both as playing a part in racial differentiation and speciation. To him museum collections and data were one of the essential tools for deciphering the course and much of the mechanism of evolution, and a sine qua non for the construction of a systematics that could aspire to an accurate presentation of taxonomical relationships. Ruthven’s twenty- three years of leadership — until 1929, when he resigned to accept the presidency of the University — were devoted to developing the Museum for this purpose and gave it its peculiar status as a university museum for research and teaching. In 1909 the geological material was transferred to the Department of Geology, leaving only the zoological and anthropological collections in the custody of the University Museum. The anthropological specimens, although accumulating and cared for, were inactive, held in trust for a future museum of anthropology. In 1913 the Regents, in view of the restricted scope of the collections and the active interest in zoological research, formally recognized the Museum of Zoology as a separate administrative unit, with Ruthven as Director. In reviewing this action in his Report for 1912-13, Ruthven stated:

This distinctly separates the museum from all other departments in the university, placing it upon the same footing 4 Museum of Zoology

as the General Library, and definitely fixes the responsibility for its development upon the person directly in charge. The museum is now in position to pursue energetically the policy that has been adopted in recent years. Briefly, this policy requires that primary attention be given to the preservation of materials for the study of the Michigan fauna, that limited explorations be made outside of the state for the purpose of acquiring material for illustration and comparison, that schools and working naturalists in the state receive such assistance as can be rendered them, and that as much research work as possible be done on local problems and in the general fields in which the members of the staff have specialized.

The staff of the newly recognized Museum of Zoology consisted of Alexander G. Ruthven, Director, Norman A. Wood, Curator of Birds, Bryant Walker, Honorary Curator of Mollusca, William W. Newcomb, Honorary Curator of Lepidoptera, Arthur S. Pearse, Honorary Curator of Crustacea, Etta Van Horn, Administrative Assistant, Crystal Thompson, Scientific Assistant in Charge of Fish and Invertebrates, Helen Thompson, Scientific Assistant in Charge of Amphibians and Reptiles, Bradshaw H. Swales, Associate in Ornithology, Arthur W. Andrews, Associate in Entomology, and Charles K. Dodge, Associate in Botany. Wood, who was to become the dean of Michigan ornithology, began his association with the Museum in 1895 as a taxidermist, employed to mount the birds of the Steere Collection, but his innate abilities and rapid development as an ornithologist and field naturalist made his appointment as Curator of Birds a particularly happy one.Helen Thompson, later Helen T. (Mrs. Frederick M.) Gaige, and Crystal Thompson had been associated with the Museum since 1911 as students of natural history and had won a place on the staff by their enthusiasm and competence. Walker, Newcomb, Pearse, Swales, Andrews, and Dodge held honorary appointments that recognized their deep interest in natural history and their enthusiastic response to the opportunity to share in a program of field and museum studies. Of these only Pearse was a professional zoologist, then at the beginning of an academic career that was to lead to professorships at Wisconsin and Duke The Museum of Zoology (1942) 5 and recognition in the development of ecology in America. Bryant Walker was a Detroit attorney who had already won as high a place in his avocation of malacology as in his profession as a corporation lawyer. Newcomb was a physician, living first in Detroit and then in Ann Arbor, with an enduring interest in the Lepidoptera; Swales, a Detroit and later a Washington, D. C., attorney, found time to carry on studies on the natural history of birds, and Andrews, a master cabinet-maker, became a serious student of the Coleoptera and for many years contributed greatly to a knowledge of the insect fauna of Michigan. Dodge, a customs collector at Port Huron, had made himself an authority on the botany of Michigan and had developed an extraordinary knowledge of plants in the field. He continued to add to knowledge of the Michigan flora until his death in 1918. This group of professionally competent amateurs who participated in the councils of the Museum took part in its program of studies and established a strong tradition of the peculiar worth of gifted and devoted amateurs in the maintenance of an esprit de corps and as associates in museum research. Walker and Newcomb had been generous in support of the earlier biological explorations carried on by Adams and Ruthven. The expeditions to Isle Royale and the Porcupine Mountains in 1904 and 1905 and the field work in 1907 in Iowa by Ruthven and Max Peet, in Dickinson County in 1909 and in Veracruz, Mexico, in 1910 by Ruthven and H. B. Baker were made possible by their financial support. These expeditions by Adams and Ruthven, the 1908 state-supported field work in Huron County, and the Mershon expeditions to the Charity Islands in Lake Huron in 1910-11 had an essential part in the emergence of the Museum of Zoology. Its reorganization as a separate department was a stimulus to increased activity. Thanks very largely to the continued generous support of Walker, Newcomb, and Swales, field work in Michigan and in regions outside the state was carried out on an extensive scale. Expeditions were sent to the Maggie Basin in Nevada in 1912, to southern Illinois and to the Santa Marta Mountains of Colombia in 1913, to British Guiana and the Davis Mountains of Texas in 1914, to Schoolcraft County in Michigan 6 Museum of Zoology

in 1915, to the Davis Mountains again and to the Appalachians of North Carolina in 1916. George Shiras, of Marquette, provided for museum field work at Whitefish Point on Lake Superior in Chippewa County in 1912, 1913, and 1916, in Alger County in 1916, and, with Bryant Walker, supported field parties in Berrien County in 1917. In 1913, through the support of Walker, Swales, and Newcomb, the Occasional Papers, the Museum’s first scientific series, was begun. In the same year Frederick Mahon Gaige (’14) was added to the staff as Scientific Assistant in Charge of Insects, and in 1914 George R. LaRue of the Department of Zoology was made Honorary Curator of Parasitic Worms. In 1916, with the support of Walker, Swales, and Newcomb, the second scientific series of the Museum, the Miscellaneous Publications, was established. This year is also memorable for the appointment of Edward Bruce Williamson (Ohio State ’95) as Honorary Curator of Odonata. Williamson, a banker and horticulturist of Bluffton, Indiana, was a student of the dragonflies. Like Bryant Walker he was an amateur who had achieved an international reputation as an authority in his field, and, like Walker, he contributed greatly to the Museum’s growing scientific eputation.r In 1916-17 much progress was made toward the present organization of the Museum into divisions: a Division of Birds and Mammals with Wood as Curator and Swales as Honorary Associate Curator in Ornithology; a Division of Reptiles and Amphibians with Ruthven as Curator and Crystal Thompson and Helen T. Gaige as scientific assistants; a Division of Insects with Frederick M. Gaige as Curator, Newcomb as Honorary Curator of Lepidoptera, Williamson as Honorary Curator of Odonata, and Andrews as Honorary Associate Curator in Entomology; a Division of Mollusks with Mina Winslow as Scientific Assistant in charge and Bryant Walker as Honorary Curator; a Division of Crustaceans with Pearse as Honorary Curator; a Division of Parasitic Worms with LaRue as Honorary Curator; and a Division of Botany with Dodge as Honorary Associate Curator. Such provision as could be made for a collection of fishes was for the time assumed by the Division of Reptiles and Amphibians. The next year Mrs. Gaige was made Assistant to the Director, The Museum of Zoology (1942) 7 and three new honorary associate curators were appointed: Cecil Billington, of Detroit, in botany, Calvin Goodrich, of Toledo, in mollusks, and James Speed Rogers, of Guilford College, North Carolina, for the order Diptera. In 1919 Miss Winslow was made Curator of Mollusks, Mrs. Gaige became Assistant Curator of Reptiles and Amphibians as well as Assistant to the Director, and Lee R. Dice was brought in as Curator of Mammals. The earlier Division of Birds and Mammals was split into separate divisions of Birds and of Mammals, with Wood and Swales now responsible for the bird collection. In 1920 a Division of Fishes was created. Carl L. Hubbs, following Walter N. Koelz, took over the then modest collection of fishes that had been cared for by the Division of Reptiles and Amphibians — a collection accumulated chiefly through the work of Professor T. L. Hankinson, Professor Jacob Reighard, and Walter N. Koelz. In 1921 the Section of Botany was transferred to the newly formed University Herbarium. This left the Museum with six major divisions — Mammals, Birds, Reptiles and Amphibians, Fishes, Insects, and Mollusks, each with a full-time curator — and with two minor divisions that were essentially repositories for incidental collections of Crustacea and parasitic worms. The growth of the collections, which had been continuous since the inauguration of the early surveys by Adams in 1903, began to accelerate markedly with reorganization of the Museum of Zoology in 1913. The intensive investigation of Michigan’s fauna had continued with more emphasis on ecological relationships and correlation with the detailed physical and historical geography of the state. Each of the divisions was also studying the distribution and ecological relationships of particular groups or faunas and sending field parties on expeditions into South America, Central America, Mexico, the mountain and desert regions of the western United States, such peculiarly isolated faunal areas as the Davis Mountains of Texas and the Olympics of Washington, and present and former river drainage systems of North America. Much work was also done by staff members in collaboration with special biological surveys of other states, and all of this 8 Museum of Zoology

added to the growth of the collections and to their increasing adequacy for research and teaching. By 1920 the old Museum Building was outgrown and an overflow into other quarters began. Several rooms in the Natural Science Building were made available for the Division of Fishes, but these were soon inadequate, and “annexes” helped to meet the increasing need for space — part of the third floor of the Old Medical Building and a succession of frame houses acquired by the University in plans for campus expansion. In 1925 the legislature appropriated funds for a University Museums Building. This was not ready for occupancy until 1928, but the congestion in the old Museum Building and its annexes was made more endurable in the knowledge that adequate quarters were in prospect. Both Adams and Ruthven thought of collections as essential tools for the investigation of an important group of broadly related problems: the bearing of geography and geological history on present and past distribution of animal species and faunas; the interaction in nature between animal populations and the kind and availability of habitats suitable for their needs. This was an aspect of natural history of common importance to many fields of theoretical and applied biology, but badly limited by the dearth of truly extensive and sufficiently documented collections of actual specimens. Because of the multiplicity of animal forms and their intricate patterns of distribution and interrelationship, the amassing of adequate collections can at best be only a gradual process, and, because of its complexity, can proceed only by alternating the study of collections at hand with fresh excursions afield to fill discovered gaps in collections and data. This concept loomed large in maintaining the support that had come from Walker, Williamson, and other honorary curators and had long been one of Ruthven’s basic teachings. His essays on “Geography in Museums of Zoology,” “Systematic Zoology in Museums,” “The Relation of the Museum to the High Schools and Grade Schools of the State,” and “Some Considerations Pertinent to the Development of a Museum Policy,” in his Report of the Director of the Museum of Zoology The Museum of Zoology (1942) 9 for the years 1920-26, attest this broadly conceived idea of the role of a University museum. The years 1925-28 were especially busy ones. In addition to the nearly normal programs of expeditions, research, and curatorial work, the translating of long-developed ideas of specimen ranges, live rooms, aquaria, divisional libraries, and laboratory-offices into working plans, and the designing of the special equipment and of the uniquely accessible shelving for the huge reptile and amphibian and fish collections were completed, and 1928 brought the huge task of installing the collections in their new quarters. The new building, which included space for the University Herbarium, the Museum of Anthropology, the Museum of Paleontology, and a Section of Exhibits, as well as for the Museum of Zoology, was a notable example of pleasing highly functional museum planning and construction. The needs of the growing Museum of Zoology for the ensuing twenty-five years were, on the whole, admirably anticipated, and only within the past few years has the Museum again begun to feel the limitations of increasingly crowded quarters. The fiscal year 1927-28 was marked by a number of important changes in staff. Ruthven was appointed to the newly created post of Director of University Museums, Gaige was made Assistant Director of the Museum of Zoology and Mrs. Gaige Assistant to the Director of University Museums. All of these appointments were in addition to those previously held. New personnel included Josselyn Van Tyne, Assistant Curator, Bird Division, Grace Eager, Museum Artist, William G. Fargo, of Jackson, Honorary Curator of Birds, and William P. Harris, of Grosse Pointe, Honorary Associate Curator of Mammals. The year also brought the loss by death of Bradshaw H. Swales, Honorary Associate Curator of Birds since 1912. The next year was Ruthven’s last as Director of the Museum of Zoology. His appointment to the directorship of the University Museums had been a part of the long-planned organization of natural history museums in the University, and had been conceived as an extension of his extraordinarily fruitful development and administration of the Museum of Zoology; but in the same month ( June, 1928) in which the new Museum 10 Museum of Zoology

was formally dedicated the Regents asked him to act as Dean of Administration for the University. In June, 1929, following the resignation of President Little, the Regents requested that he “divest himself so far as possible, for the present, of his responsibilities centering about the Museum and the Department of Zoology” and that he “allow his resignation as Dean of Administration, which he had desired to have the Board consider, to lie upon the table” (R.P., 1926-29, p. 1017). Then, in October of 1929, he was asked to accept the presidency of the University. Ruthven hoped to keep his close association with the Museum, and he retained his positions, Director of University Museums and Curator of Reptiles in the Museum of Zoology, as long as there was any hope that the duties of the presidency could be lightened sufficiently to allow a part-time return to herpetological research and museum policies. Instead, he was soon faced with the problems of University administration in a great depression and then in World War II. The problem of filling the directorship of the Museum of Zoology was chiefly one of persuading Gaige to accept administrative duties at the expense of his research and concern with the Insect Division. He had been associated with the Museum since 1908, and had been a member of many of its expeditions. He had been Curator of Insects since 1916 and had not only assisted Ruthven in the development of the Museum, but his extraordinarily broad knowledge of natural history, its literature and workers, and his unselfish loyalty to the Museum made his appointment in November, 1929, a happy one. Gaige’s first Report of the Director of the Museum of Zoology for the year ending June, 1929, issued early in 1930, lists the following staff:

• Alexander G. Ruthven, Director of University Museums • Frederick M. Gaige, Director of Museum of Zoology • Helen T. Gaige, Assistant to Director of University Museums • Geneva Smithe, Secretary of University Museums • Crystal Thompson, Curator of Department of Visual Education The Museum of Zoology (1942) 11

• Kimber C. Kuster, Librarian • Morley W. Williams, Superintendent of Building • Scientific taffS

◦ Lee R. Dice, Curator ◦ William P. Harris, Jr., Associate Curator ◦ Adolph Murie, Assistant Curator ◦ Ruth D. Svihla, Assistant

• Division of Birds

◦ Norman A. Wood, Curator ◦ Josselyn Van Tyne, Assistant Curator ◦ William G. Fargo, Honorary Curator ◦ Walter E. Hastings, Custodian of Birds’ Eggs

• Division of Reptiles and Amphibians

◦ Alexander G. Ruthven, Curator of Reptiles ◦ Helen Thompson Gaige, Curator of Amphibians ◦ Howard A. Kelly, Honorary Curator ◦ Norman E. Hartweg, Assistant

• Division of Fishes

◦ Carl L. Hubbs, Curator ◦ Walter Koelz, Assistant Curator ◦ Laura C. Hubbs, Cataloguer

• Division of Insects

◦ Frederick M. Gaige, Curator ◦ E. B. Williamson, Research Associate ◦ Samuel A. Graham, Research Associate ◦ William W. Newcomb, Honorary Curator of Lepidoptera ◦ Sherman Moore, Associate Curator of Lepidoptera 12 Museum of Zoology

◦ James Speed Rogers, Associate Curator of Diptera ◦ Arthur W. Andrews, Associate Curator of Coleoptera ◦ Ada Olson, Assistant

• Division of Mollusks

◦ Mina L. Winslow, Curator ◦ Calvin Goodrich, Assistant Curator ◦ Bryant Walker, Honorary Curator ◦ Ruth Morris, Assistant

• Division of Crustaceans

◦ Edwin P. Creaser, Assistant in Charge

• Division of Annelids

◦ Frank Smith, Honorary Curator

• Division of Parasitic Worms

◦ George R. LaRue, Research Associate

• Division of Protozoans

◦ Dora S. Lemon, Custodian

• Division of Extension

◦ Crystal Thompson, Curator

• Technical Staff

◦ Carleton W. Angell, Sculptor, University Museums ◦ Grace Eager, Artist ◦ James Wood, Preparator ◦ A. Russell Powell, in charge of shop ◦ Elsa Hertz, Telephone and Information Clerk

Of the eleven divisions listed in the report, the first six — Mammals, Birds, Reptiles and Amphibians, Fishes, Insects, and The Museum of Zoology (1942) 13

Mollusks — were strongly entrenched with extensive and rapidly growing research collections, full-time staffs, and expanding research and teaching programs. With the exception of the Fish Division their origins had antedated the organization of the Museum into divisions. The divisions of Crustaceans, Annelids, Parasitic Worms, and Protozoans were set up as tentative trials of the feasibility of establishing such collections without continued curatorial care, solely with the interested collaboration of honorary curators. The Division of Crustaceans owed its origin to Professor A. S. Pearse of the Zoology Department, an interest and collaboration that continued for several years after Pearse left Michigan. In 1929 the division was reactivated with the appointment of Edwin P. Creaser, a graduate student of crayfishes, as Assistant in Charge. He was promoted to Curator in 1933 and resigned in the same year. No further curatorial appointments were made. During and following Creaser’s curatorial activity the crustacean collection grew considerably. It remained inactive, however, and in 1950 the Museum obtained permission of the Regents to present it to the National Museum, where it is assured of continued accessibility to workers in this group. The divisions of Annelids, Parasitic Worms, and Protozoa had an even briefer existence than that of Crustaceans, and none survived the interest and collaboration of their honorary curators or custodians. The collection of parasitic worms built up by Professor George R. LaRue and his students, of the Department of Zoology, was transferred to the Bureau of Parasitology, Agricultural Research Center, Beltsville, Maryland. This was done at LaRue’s request when, in 1949, he retired and moved to Beltsville to continue his studies in helminthology. The Division of Extension was later combined with the Department of Visual Education and transferred to the University Museums. Both extension and visual education were under the direction of Crystal Thompson and formed the basis for the present Section of Exhibits of the University Museums. The year 1930 was notable for the gift of the Edwin S. George Reserve to the University. The Reserve, a tract of approximately two square miles of rolling knob-and-basin topography with 14 Museum of Zoology

extensive areas of woods, old fields, marshes, and swamps, providing a wide range of animal and plant habitats of southern Michigan, was accepted as a native wildlife preserve to be administered by the Museum of Zoology. It has proved a magnificent laboratory in natural history not only for the study of an abundant natural biota but as an area where field studies may be carried on safe from the interference of fires, tillage, trespass, or change of ownership. Gaige’s directorship extended from 1929 until August, 1945, through the difficult depression years of the 1930’s and World War II. Although he never lost his distaste for the routine forms and minutiae of administration he carried on as Director through the academic disruption of the war years, but a few weeks after the final surrender of Japan he resigned both the directorship and the curatorship of the Insect Division. Mrs. Gaige resigned as Curator of Amphibians on the same date, and the double loss was severely felt by the entire Museum. Gaige’s outstanding contributions as Director had been his loyal support of his staff and the example of his extraordinarily broad knowledge of natural history. The friendship and moral and material support that he and Mrs. Gaige gave to so many beginning and often uncertain younger naturalists became a tradition to a whole generation of museum students. William H. Burt, Curator of Mammals and chairman of the Museum’s Executive Committee, served as administrative head from the time of Gaige’s resignation until 1947, when James Speed Rogers, (’15, Ph.D. ’31), Honorary Associate Curator of Diptera since 1918, a former student of both Ruthven and Gaige, and Professor of Biology at the University of Florida since 1922, was appointed Director. The account of the Museum since 1929 can best be related in a brief review of the divisions. Mammals. — The establishment of a separate division for mammals dates from 1919, when Dr. Lee Raymond Dice (Stanford ’11, Ph.D. California ’15) was appointed Mammalogist and took over the mammal collection that until then had been the responsibility of the Curator of Birds. The first objectives set by Dice were the rounding out of the modest and rather random collection and intensive field work on the mammals of The Museum of Zoology (1942) 15

Michigan. This was soon supplemented by field work in other areas pertinent to the needs of the collection and to the interests of the curator, and special attention was given to the hares, rabbits, and coneys. In 1927 the enthusiastic co-operation of William P. Harris, of Grosse Pointe, was recognized by his appointment as Honorary Associate Curator, a relationship that continued to be a most happy one for the Museum. His studies on the North American squirrels and the generous sponsorship he has given through the Harris Fellowship in Mammalogy, the purchase of specimens, and the support of expeditions have greatly aided the division. In 1931 Philip M. Blossom was appointed Honorary Associate Curator. He contributed materially to the stature of the Museum, particularly in his work in the Southwest on desert mammals. Dice became particularly interested in the bearing of genetics and ecological factors on the evolutionary relationships of wild mammal populations and increasingly preoccupied with his own important researches in these fields. Breeding stocks of wild rodents were established to test the inheritance of characters used in systematics, and a long series of breeding experiments was planned and carried out on crosses between taxonomically distinct races and species. This work was greatly furthered by the gift of the breeding stocks studied by Sumner at the University of California at La Jolla, and by close co- operation with the Laboratory of Mammalian Genetics. In 1929 Adolph Murie (Concordia ’25, Ph.D. Michigan ’29), who had taken his doctorate in mammalogy under Dice, was appointed Assistant Curator and took over much of the responsibility for the general mammal collection. He was succeeded by Seth B. Benson in 1934-35, and Benson by William Henry Burt (Kansas ’26, Ph.D. California ’30) in 1935. In 1934 Dice was also made Director of the Laboratory of Vertebrate Genetics, and in 1938 he resigned as Curator of Mammals to direct the Laboratory of Vertebrate Genetics. Burt was made Curator of Mammals, and Dice became Honorary Associate Curator, a position he continued to hold until 1946. In 1938 Emmet Thurman Hooper (California ’33, Ph.D. ’39) was appointed Assistant Curator of Mammals, becoming 16 Museum of Zoology

Associate Curator in 1946. Under Burt and Hooper the division has continued to expand and to maintain an active program. Burt’s own researches and those of his students have been chiefly in the ecology of wild populations, especially the behaviors associated with home ranges and other manifestations of territoriality. Hooper’s studies have centered chiefly about the derivation and relationships of the mammal fauna of Mexico, particularly the geographic factors in speciation and evolution. Much of Burt’s and Hooper’s concern has been to add to the knowledge and appreciation of Michigan’s mammal fauna and resources, a project that has been popularly summarized in Burt’s The Mammals of Michigan. Birds. — The bird collection had been the special concern of Norman A. Wood in the days of the old Museum when Wood was employed as a taxidermist for the birds of the Steere Collection. Adams recognized Wood’s ability and intimate knowledge of birds and gave him the responsibility for ornithology in the expeditions to the Porcupine Mountains and Isle Royale. Under Ruthven, Wood carried on studies of migration at Point Pelee, Ontario, and continued to be responsible for both ornithological field work and the care of the bird collection, a status that was officially recognized by his appointment as Curator in 1911. The division was also fortunate in the support it continued to receive from honorary curators and other collaborators. Bradshaw H. Swales was Honorary Curator from 1912 until his death in 1928, Walter E. Hastings, Custodian of Birds’ Eggs from 1921-32, and William G. Fargo, Honorary Curator from 1927 to 1943. Fargo, a civil engineer, has since 1926 given much support to the division by his own field work, the purchase of extensive collections, and generous financial aid for expeditions. Other longtime collaborators and generous sponsors were Dr. Max M. Peet of the University’s surgical staff, an enthusiastic student and collector of birds since his early student days, A. D. Tinker, of Ann Arbor, and the Michigan Department of Conservation. Josselyn Van Tyne (Harvard ’25, Ph.D. Michigan ’28) was appointed Assistant Curator in 1928 and Curator in 1931; in 1933 Wood, after some forty years of continuous studies, chiefly on The Museum of Zoology (1942) 17 the birds of Michigan, was made Curator Emeritus. William Pierce Brodkorb (Illinois ’33, Ph.D. Michigan ’36) was appointed Assistant Curator in 1936, a position he held until 1946. He was succeeded by Joseph James Hickey (New York Univ. ’30, Ph.D. Michigan ’49). After Hickey’s resignation George Miksch Sutton (Bethany College, ’23, Ph.D. Cornell ’32) was appointed Curator, half time, reserving the remainder of his time for his bird paintings and illustrations. In 1949 Sutton resigned the curatorship to devote himself to his illustrations and research but remained as Research Consultant until he resigned in 1952 to accept a professorship at the University of Oklahoma. In 1949 Robert W. Storer (Princeton ’36, Ph.D. California ’49) was appointed Assistant Curator. The collections of the Bird Division have been guided largely by three chief objectives: the development and maintenance of an exhaustive and precisely documented representation of Michigan and North American birds, an adequate synoptic collection of the families and genera of the birds of the world, and the establishment of strong research collections in those groups of especial interest to the staff. Much attention is also given to assembling adequate skeletal and other anatomical series needed for the critical revision of familial and intergeneric relationships. Even so brief a review of the Bird Division would be remiss if it failed to stress the indefatigable work of Van Tyne in building up the increasingly adequate research collection and the outstanding ornithological library that so greatly facilitates the work of the division. An exceptional series of gifts has contributed greatly to the growth of the division. Among the more notable gifts have been the Walter Koelz collection of 3,350 specimens, one of the many important gifts of William Fargo; the huge collection of Dr. Max M. Peet, extraordinarily rich in many rare species and series, the gift of Mrs. Max M. Peet; the Shufeldt collection of Mexican birds; and the extensive collections made in Costa Rica by Paul Slud. Reptiles and Amphibians. — The cold-blooded vertebrates had received scant attention in the Museum prior to the Isle Royale expeditions of 1904 and 1905, when Ruthven began an intensive study of Michigan herpetology. By 1910 he had the 18 Museum of Zoology

enthusiastic collaboration of Helen Thompson (later Mrs. F. M. Gaige) and Crystal Thompson. They had accumulated sufficient collections and data by 1912 to publish the first Herpetology of Michigan, in which forty-four species of reptiles and eighteen of amphibians were recorded from the state. Although the concern with Michigan fauna continued to be a primary interest of the division, explorations were carried much farther afield. The Maggie Basin of Nevada was visited in 1912, and in 1913 the expedition to the Santa Marta Mountains of Colombia initiated the series of trips to northern South America, Central America, and Mexico that were to give the Museum notable collections of a rich neotropical fauna on which Ruthven, Mrs. Gaige, and their students were to base a long series of important papers. By 1917, when the organization of the Museum into divisions was begun, the reptile and amphibian collections were already extensive and important, and the Museum had acquired a reputation as a strong center of herpetological research. In 1918 Mrs. Gaige was made Assistant Curator, and in the same year Frank N. Blanchard began a doctoral problem under Ruthven on the kingsnakes of the genus Lampropeltis. This was the beginning of a program of graduate teaching and research in which a large proportion of the American workers in herpetology of the present generation were trained. The excellence and efficiency of this training, and Ruthven’s ability to keep in touch with it so long after he had become President of the University, owed much to the long and close co-operation between Ruthven and Mrs. Gaige. In 1927 Ruthven became Curator of Reptiles and Mrs. Gaige Curator of Amphibians, an arrangement that recognized Mrs. Gaige’s share in the work and research of the division. Within a little more than a year, however, it was necessary for Mrs. Gaige to assume curatorial care of the entire division and to take increasing responsibility for much of the direction of graduate instruction. Norman Hartweg (’30, Ph.D. ’34) was added to the staff as Assistant Curator in 1934 and became Associate Curator in 1942. In 1945, after Mrs. Gaige’s resignation, Hartweg was given charge of the division and was made Curator in 1946. In 1947 Charles Frederic Walker (Ohio State ’30, Ph.D. Michigan ’35), formerly The Museum of Zoology (1942) 19 associate professor at Ohio State University, became Associate Curator. Both Hartweg and Walker obtained their doctorates under Ruthven and Mrs. Gaige, and they have continued the old traditions as far as has been practicable with an expanding teaching program and an ever-growing collection. Hartweg’s primary research interests have been largely with turtles, Walker’s with the amphibians, but their students’ researches have included almost the entire taxonomic range of herpetology. Fishes. — The small fish collection accumulated over the years was augmented in 1919 by the gift from the U. S. Fish Commission of some 3,000 specimens from the Great Lakes region, consisting largely of whitefishes collected by Walter Norman Koelz (Olivet ’15, Ph.D. Michigan ’20) in furtherance of his monographic study of that group. Koelz was appointed Curator of Fishes by the Regents in December, 1919, but left the following March to take a position with the U. S. Bureau of Fisheries. Carl Leavitt Hubbs (Stanford ’16, Ph.D. Michigan ’27), a student of Charles Henry Gilbert and protégé of David Starr Jordan at Stanford, assumed the curatorship in July, 1920, and the present Division of Fishes, with its unrivaled collection of North American freshwater fishes and excellent synoptic representation of other faunas, is largely a monument to his untiring energy, enthusiasm, and ability. There was no room for another collection in the overcrowded old Museum Building. Instead, temporary and often makeshift quarters were found. Two rooms provided in the new Natural Science Building in 1919 were outgrown by 1921, and the collections were moved to the Museum Annex, a frame house on East University Avenue, then into rooms in the old Medical Building, and once more into the Natural Science Building. In spite of handicaps the collections grew rapidly. The first concern was to build up the Michigan collection, and the first two years were devoted to this objective. Of the more than 50,000 specimens accessioned for 1922-23, some 18,000 were taken by Hubbs in California, a large collection was obtained in North Dakota by T. L. Hankinson, Professor at the Michigan State Normal College and a long- 20 Museum of Zoology

time collaborator, and approximately 20,000 specimens from Michigan or the Great Lakes were received through the co- operation of the United States Bureau of Fisheries and by the field work of Hubbs and his students. Hubbs had remarkable success in obtaining active co-operation from other individuals and institutions, in large part because of the essential aid he provided in making taxonomic determinations and the sound practical suggestions which he was able and willing to give to workers on fishery problems and surveys. As a consequence the division became a repository for many of the collections made by state surveys, the Great Lakes section of the United States Bureau of Fisheries, and many independent workers. The co-operation with the Michigan Department of Conservation and the Great Lakes Division of the Bureau of Fisheries has continued to be especially close and important. In 1925 Jan Metzelaar (Sc.D. Amsterdam ’19), fishery expert for the Michigan Department of Conservation, was made Honorary Custodian of Michigan Fishes, in recognition of his work with the Fish Division since 1923. This continued until his untimely death, by drowning, in 1929 while engaged in fishery investigations. In 1927 Mrs Laura C. Hubbs received appointment as Cataloguer of Fishes, in view of her work as a volunteer, and in 1929 Walter Koelz returned as Assistant Curator of Fishes. Hubbs organized the Institute for Fisheries Research in 1930 under the sponsorship and with the support of the Michigan Department of Conservation and was its director until 1935, as well as Curator of the Fish Division. Koelz resigned as Assistant Curator to accept the post of Ichthyologist in the Institute, and Carroll Willard Greene (’25, Ph.D. ’34) was made Assistant Curator. Hubbs’s broad interest in the evolution and systematics of fish had brought him in contact with the problem of hybridization in nature, a question of great theoretical importance in systematics and evolution and one that is particularly pertinent and pressing in ichthyology. In 1927 he began a series of breeding experiments to test the possibilities, limits, and results of crossing taxonomically distinct forms, experiments that were soon expanded to the capacity of the aquarium room in the The Museum of Zoology (1942) 21 new building. The long series of aquarium investigations, in which Mrs. Hubbs and his graduate students were important collaborators, was supplemented by extensive and detailed analyses of hybridization in nature, based upon studies of many extensive series and of ecological and geographic factors. These investigations, continued until 1944, resulted in a number of publications of marked importance in general systematics and evolutionary theory as well as in fish taxonomy. The studies on hybridization did not lessen the other activities of the division. The collections continued to expand with attention both to taxonomic representation and to the correlation of distribution with present and former drainage systems. Staff and student publications on this accumulating material appeared as reports on regional faunas and on ecological and geographic distribution, as descriptions of new forms and groups, and as taxonomic revisions of genera and families. In 1930 Greene resigned as Assistant Curator and was replaced by John Greeley (Cornell ’25, Ph.D. ibid. ’30), until 1934, when he became ichthyologist for the New York Department of Conservation. He was replaced by Miltion Bernhard Trautman, who resigned in 1939 to accept a post with the Franz Theodore Stone Laboratory at Put-in-Bay, Ohio. The position of Assistant Curator was not continued after Trautman’s resignation; instead, provision was increased for graduate assistantships. Hubbs resigned in 1944 and accepted a professorship of biology at the Scripps Institution of Oceanography at La Jolla, California, having served twenty-four years as Curator of Fishes. In that time the collections had grown from 5,000 to almost 2,100,000 specimens, the Museum had become the outstanding center for the systematics of North American fishes, and more than forty graduate students had received training in ichthyology or fishery biology. Reeve Maclaren Bailey (’33, Ph.D. ’38), who had been assistant professor at Iowa State College and head of the Iowa Fisheries Research Unit, was appointed Associate Curator to replace Hubbs in 1944, and William Alonzo Gosline III (Harvard ’38, Ph.D. Stanford ’41) was added to the staff as Assistant Curator in 1945. Bailey was promoted to Curator in 1948, and, also in 1948, 22 Museum of Zoology

when Gosline resigned to accept a professorship in zoology at the University of Hawaii, Robert Rush Miller (California, ’38, Ph.D. Michigan ’44), formerly associate curator at the National Museum, was appointed Associate Curator. Both Bailey and Miller were trained in the Hubbs tradition, having completed their doctorates under his direction. They have continued the program of staff and student research based upon the rich store of material and have carried on extensive field explorations to fill gaps in geographic, ecological, and taxonomic coverage. Insects. — The organization of this collection dates from 1913, when Frederick M. Gaige became the Assistant in Charge of Insects. Two honorary appointments had already been made in entomology: Dr. William W. Newcomb, Honorary Curator of Lepidoptera in 1909, and Arthur W. Andrews, Associate in Entomology in 1912. Both Newcomb and Andrews were enthusiastic students of Michigan insects and were eager to help build up the Museum’s collections as soon as the specimens could be assured of proper care. Aside from a small series of Michigan moths, butterflies, and beetles contributed by Newcomb, Andrews, and other members of the Detroit Natural History Club, there were a few incidental collections of insects made by earlier Museum field parties, some remnants of the old Beal-Steere collections of the 1870’s, an extensive collection of Philippine insects presented to the Museum by Professor E. M. Ledyard of the University of the Philippines, and the ants taken by Gaige in the 1912 expeditions to the Charity Islands in Lake Huron and the Maggie Basin in Nevada. In 1913, however, two immediate objectives were established. One was an adequate representation of the Michigan insect fauna, the other a comprehensive collection of the ants, the group which Gaige had chosen for his own research. As a member of the 1913 and 1914 expeditions to Colombia and British Guiana, respectively, Gaige began the Museum’s extensive collections of neotropical ants. The Newcomb expedition to the Davis Mountains of Texas in 1914 paid especial attention to securing ants, Lepidoptera, and beetles, and in 1915 the field party sent to Schoolcraft County in the Upper The Museum of Zoology (1942) 23

Peninsula included three entomologists in addition to Gaige and Andrews. In 1916 a Division of Insects was created with Gaige as Curator, and E. B. Williamson was appointed Honorary Curator of Odonata. Williamson was a well-known amateur authority on the Odonata; he had already given much help in the determination of the dragonfly material and shared the Museum’s great interest in the neotropical fauna. The general pattern for the development of the division was beginning to take a definite form. The ants and dragonflies were to be studied on a world-wide basis with especial attention to North America and the neotropical region. For other insects the special concern was to be the Michigan fauna. Both goals were ambitious ones and were necessarily planned as accumulative projects. The most rapid progress on the Michigan fauna was in the butterflies and moths, beetles, and various conspicuous families of other orders, such as the cicadas, bumblebees, robberflies, and horseflies, in which the cordial co-operation of outside specialists such as W. T. Davis, T. H. Frison, and J. S. Hine was available. The accession of specimens was not, of course, limited to the above groups. Insects of nearly all orders were collected and preserved with the proper field data, to be stored unworked for the day when a competent investigator would undertake their study, and several such workers were in time found or trained. In 1918 J. Speed Rogers, who as a member of the Schoolcraft expedition of 1915 had begun the study of the craneflies, was made Honorary Associate Curator of Diptera. T. H. Hubbell, as a student Aid in Entomology, 1919-22, developed an enduring interest in the Orthoptera that was to result eventually in the Museum’s outstanding research collection in that group. At about that time, Roland F. Hussey, a student in the Department of Zoology, began work on the Hemiptera and as a member of several field expeditions and by much collecting about Ann Arbor developed a very respectable representation of the Michigan Hemiptera. Melville T. Hatch, another student of entomology in the Department of Zoology and a specialist in the Coleoptera, collected intensively about Ann Arbor and in Cheboygan County and gave much aid in the determination and arrangement of the beetle collection. 24 Museum of Zoology

The rapid growth of the division brought difficulties. Despite the small size of an individual insect the material was beginning to overflow all available space in the old Museum. Fortunately, several of the honorary curators were able to provide quarters for their rapidly growing accessions until the new museum building became available, and for several years perhaps less than half of the annual accessions were actually deposited in the Museum. In 1926 Sherman Moore, an engineer of the United States Lake Survey, was made Honorary Associate Curator of Lepidoptera. He had already given much attention to the collection and to the study of moths and butterflies of the Great Lakes region, and eventually (1952) was to complete the critical determination and arrangement of the extensive collection of Michigan Lepidoptera. The year 1929-30 brought many important changes, one of them unforeseen and unplanned. Gaige was made Director of the Museum of Zoology while still holding his curatorship, an increasing responsibility that gradually brought an end to his work on ants. E. B. Williamson, who had retired from his profession as a banker, accepted a full-time position as Research Associate in Odonata, and Professor Samuel A. Graham was appointed honorary Research Associate. The years 1928-33 were notable for the tremendous development of the Odonata collections and active research on this group. Williamson’s own private collection, perhaps the most extensive in North America, was given to the Museum, and the combined collections were further expanded by an energetic program of field work. Leonora K. Gloyd was made Assistant in Odonata, and in 1932 Justin W. Leonard began his doctoral problem under Williamson’s direction on a group of the neotropical dragonflies. Williamson’s death in 1933 was a severe loss to the division. Mrs. Gloyd, who continued to care for the collection until 1938, completed one of Williamson’s unfinished papers and carried on a number of studies of her own. Leonard completed his doctoral dissertation in 1937 under Gaige’s direction. In 1935 T. H. Hubbell was made Honorary Associate Curator of Orthoptera in recognition of his already important research The Museum of Zoology (1942) 25 and the extensive collection of North American Orthoptera he was establishing in the Museum. From the mid-thirties until 1946 the division was concerned chiefly with the development of the Michigan collections. Particularly intensive studies were carried on at the Edwin S. George Reserve, with Sherman Moore, Wilbur MacAlpine, W. F. Lawler, G. W. Rawson, W. C. Stinson, and J. H. Newman, working on Lepidoptera, Andrews on Coleoptera, Irving J. Cantrall on the Orthoptera, Rogers on the craneflies, and George Steyskal on the higher Diptera. All of these men except Cantrall and Rogers were members of the Detroit Natural History Club or its successor, the Detroit Entomological Club, and competent amateur specialists in the groups they studied. Irving James Cantrall (’35, Ph.D. ’40), a graduate student who was engaged on a doctoral problem on the ecology of the Orthoptera under Hubbell, in 1949 became Curator of the Reserve. The extensive collections of Orthoptera and of craneflies grew rapidly, but were to a large extent either in the laboratories of their nonresident honorary curators or in compact storage in the Insect Division. From the date of Gaige’s resignation in 1945 until January 1, 1947, the division was in the charge and under the efficient care of Ada L. Olson, Senior Technical Assistant in Entomology, who maintained cooperation with the honorary curators and other collaborators. In July, 1946, Professor Theodore Huntington Hubbell (’21, Ph.D. ’34), of the Department of Biology of the University of Florida, was appointed Curator of Insects, effective January 1, 1947, and under his direction marked progress has been made in reorganizing the collections, determining a great backlog of unworked material through the aid of specialists, and meeting the problems of expanding collections and limited storage facilities. J. Speed Rogers, in his research and teaching, since 1947 has been a member of the division and has shared with the curator responsibility for active resumption of staff and student research on insects. Although emphasis is on the Orthoptera and craneflies, graduate students are carrying on studies in ants and dragonflies. A system of summer curatorships has been established, whereby recognized specialists can be brought to the University to study and arrange the Museum’s collections 26 Museum of Zoology

in insect groups outside the competence of the resident or honorary staff. This makes authoritatively determined specimens available for reference and teaching and leads to the publication of research papers based on the Museum’s collections. Dr. F. N. Young of Indiana University has spent two summers on the water beetles, Dr. Roland F. Hussey of the University of Florida two summers on the Hemiptera, and Mrs. Leonora K. Gloyd of the Illinois Natural History Survey a summer on the Odonata. The collection of Odonata again has been greatly augmented. In 1951-52, the Museum acquired the huge collection of dragonflies and the magnificent Odonata library of Dr. Clarence H. Kennedy of Ohio State University. The combined Williamson and Kennedy collections form an unrivaled archive of dragonfly taxonomy and distributional data and, though at present relatively inactive, constitute the chief of the Museum’s outstanding research series of insects, the others being the Orthoptera, craneflies, and ants. Mollusks. — A mollusk collection had existed since the Beal- Steere expedition of the 1870’s but, except for a few gifts of small and miscellaneous lots of specimens, was dormant until 1903. Then, with the inauguration of a vigorous program of field work by Adams, a steady influx of material began. The renascent Museum soon attracted a series of gifts, notably from and through Bryant Walker of Detroit, an amateur student of mollusks who had gained an international reputation as an authority on the North American land and fresh-water fauna. In 1909 Walker was made Honorary Curator of Mollusca and in 1912 was given an honorary degree in recognition of his outstanding contribution to a knowledge of the classification and distribution of North American mollusks. Mina Winslow (Smith ’13, M.A. Michigan ’16) was added to the staff as Scientific Assistant in 1916, and the formal organization of the collections was begun. Another skilled amateur student, Calvin Goodrich of Toledo, Ohio, in 1917 was made Honorary Associate Curator. A Division of Mollusks was set up in 1918 with Miss Winslow as Curator, and through the generosity of Mr. Goodrich, Miss Doreen Potter was employed as a student assistant in the division. Policies for the development of the division included the The Museum of Zoology (1942) 27 amassing of a collection of Michigan mollusks with attention to local and ecological distribution, the development of a synoptic representation of the mollusk fauna of the world, and the accumulation of ample series in the groups selected for intensive study. Both Walker and Goodrich had an important part in the researches of the division and produced a notable series of important papers on the classification and distribution of North American mollusks. Miss Winslow was largely concerned with the Michigan fauna, and, in addition to much important work on faunal lists and bibliographies, she added materially to a knowledge of the state’s fauna and its distribution. She also gave attention to the synoptic collection of the world fauna and by visits to European museums and an expedition to South Africa arranged many important exchanges and made extensive additions to the collection. Miss Winslow resigned in 1929, and Goodrich, who had recently retired from newspaper work, was made Curator; Henry van der Schalie, then a graduate student in malacology, was appointed Assistant. That year also brought an end to Walker’s long collaboration in the researches of the division and to his even longer career as a malacologist. Goodrich’s first report as Curator of Mollusks (in the Report of the Director for 1929-1930) ends with the following paragraph:

Because of illness, Dr. Bryant Walker — for the only twelve months in forty-one years — has been unable to add to his writings upon mollusca. Dr. Walker’s first paper, written in collaboration with Mr. C. E. Beecher, appeared in the Proceedings of the Ann Arbor Scientific Association for 1875-76, and in 1879 he published the first of his several catalogues of Michigan mollusks, but it was not until 1889 that he began upon the scarce interrupted begetting of papers.

Walker never fully regained his health and died in May, 1936. His collections and library were willed to the division: 100,000 lots of shells and 1,500 volumes. This bequest was a major factor in giving the division its outstanding position as a center for research and instruction in malacology. 28 Museum of Zoology

In 1931 Allan F. Archer began graduate work in mollusks under Goodrich and served as a voluntary assistant in the division until 1936, when he completed his doctorate and was appointed Assistant. He resigned in 1937 to accept a Rackham fellowship. In 1934 Henry van der Schalie (Calvin College ’29, Ph.D. Michigan ’34) was made Assistant Curator. On Goodrich’s retirement in 1944 with the title of Curator Emeritus, van der Schalie was made Curator. His curatorship has been marked by important changes in the division. The program of graduate teaching has been greatly expanded, and a course and a seminar in malacology have been established. He and his students have continued to develop the collection but with several important changes in emphasis. There is concern with life histories and ecology, a revival of interest in the Michigan fauna, and active collaboration with public health agencies in relation to control of snail-borne diseases. Publications. — The publications of the Museum of Zoology are in two main series — the Occasional Papers and the Miscellaneous Publications — made possible by funds donated by Walker, Swales, and Newcomb. The Occasional Papers, begun in 1913, are based principally on the Museum collections. The papers are issued to libraries and specialists. More than 570 have appeared. The Miscellaneous Publications are monographic studies and other contributions not within the scope of the Occasional Papers. More than ninety have been published. Other Museum publications are in the Michigan Handbook Series.

SELECTED BIBLIOGRAPHY

Adams, Charles C.Report of the Curator of the University Museum … Ann Arbor: Univ. Mich., 1903-6. Ark, The. Ann Arbor: Museum of Zoology, 1922-32. Gaige, Frederick M.Report of the Director of the Museum of Zoology … Ann Arbor: Univ. Mich., 1929-34. President’s Report, Univ. Mich., 1921-54. Proceedings of the Board of Regents …, 1865-1954. Rogers, J. Speed. Report of the Director of the Museum of Zoology … Ann Arbor: Univ. Mich., 1947-54. The Museum of Zoology (1942) 29

Ruthven, Alexander G.A Naturalist in a University Museum. Ann Arbor: Privately Printed, 1931. Ruthven, Alexander G.Report of the Director of the Museum of Zoology. Ann Arbor: Univ. Mich., 1913-29. Ruthven, Alexander G.Report of the Head Curator of the University Museum … Ann Arbor: Univ. Mich., 1907-12. (Title varies.) The University Museums Building of the University of Michigan. Ann Arbor: Univ. Mich., 1929. [2]

The Museum of Zoology (2015)

Diarmaid Ó Foighil

The Museum of Zoology (UMMZ) is the University of Michigan’s center for the study of animal diversity. UMMZ focuses on the evolutionary origins of the planet’s animal species, the genetic information they contain, and the communities and ecosystems they help to form. Now an integral part of the Department of Ecology and Evolutionary Biology (EEB), the UMMZ houses world-class collections that span almost 200 years of regional and global biodiversity studies and that support a multi-faceted departmental research and teaching program.

Beginnings: 1837-1913

The UMMZ’s collections date to the foundation of the state of Michigan. The 1837 initial session of the Michigan legislature authorized a State Geological Survey to compile information about the region’s geological features and to inventory its flora, fauna, and fossils. To house the inventoried specimens, it The Museum of Zoology (2015) 31 established a Cabinet of Natural History in the University of Michigan. Dr. Abram Sager headed the Botanical and Zoological Departments of the State Geological Survey. In 1839, he wrote a segment of the 2nd Geological Survey report entitled, A Systematic Catalogue of the Animals of the State, so far as Observed. The report listed the various species of animals encountered, especially birds. Sager’s specimens formed the initial nucleus of the UMMZ’s collections, and the museum retains twenty-two of his bird specimens, many of them in good condition. When the University relocated to Ann Arbor in 1837, Asa Gray was appointed professor of botany and zoology. But Gray began his duties with a book-buying tour of Europe for the future University Library, and he spent little time in Ann Arbor before resigning in 1842 to take a position at Harvard, where he enjoyed a distinguished botanical career, including active collaboration with Charles Darwin. Dr. Abram Sager was appointed as Gray’s replacement in 1842 and continued in this role until transferring to the newly organized Medical Department in 1850. From the beginning, the natural history cabinet specimens, located in Mason Hall, played a central instructional role on the Ann Arbor campus. The first classes in zoology consisted of lectures supplemented with demonstrations of cabinet specimens, many collected by Sager himself. A taxidermist was hired on a piecework basis to take care of the specimens, and by 1850 the cabinet consisted of up to 5,000 Michigan geological, zoological, and botanical specimens. Alexander Winchell, who was appointed professor of geology, zoology, and botany in 1855, expanded the zoological curriculum and the associated teaching collection beyond the Michigan fauna by acquiring, for example, the Trowbridge Collection of Pacific Coast species from the Smithsonian Institution in 1858. To transform the Cabinet into what would become the Museum of Natural History, Winchell hired Carl Rominger, M.D., as assistant curator, paleontologist, and taxidermist in 1863. By 1867, the zoological collection contained some 16,000 specimens. Winchell resigned in 1873; he was replaced in the short term by Eugene Hilgard (1873-75) and Mark Harrington (1875-76). Prior to the 1870s, the natural history collections were 32 Museum of Zoology

primarily national in scope. This changed dramatically when the University hired Joseph Beal Steere to lead a five-year circumglobal expedition (1870-1875) to collect materials in the natural and human sciences for the Museum. Upon his return in 1876, Steere was appointed assistant professor of paleontology and curator of the Museum. The Steere Zoological Collection alone totaled 60,000 specimens, including 25,000 insects, 1,500 mollusks and 8,000 birds. Steere was particularly interested in birds and made historically important collections in the Amazon, Peru, Malaysia, Taiwan, Celebes, and the Philippines. He was the first naturalist to collect in many of these places, especially the Philippines, and his bird collections contained more than 50 species new to science. The flood of new material for the collection created a severe space crunch. Many specimens were housed in less than ideal locations, including attics. The University president’s report of 1878 noted the unacceptable fire risk posed to “our rare and extensive scientific collections;” this led to funding for construction of a new University Museum Building in 1879-80. The natural history collections migrated to their new home in the fall of 1880. In 1879, zoology was separated from both botany and paleontology, and Steere became professor of zoology and curator of the University Museum. Coincident with the museum building construction, the Board of Regents approved a new set of operating rules stating that professors in charge of instruction in the various fields of natural history be “the curators of the corresponding collections in the Museum of Natural History.” One long-term problem with this decentralized arrangement was that collections developed unevenly; they reflected the individual interests of faculty, and the zoological collections grew faster than the others. Jacob Reighard joined the faculty in 1886, and instruction in zoology was divided into general zoology, taught by Steere, and animal morphology, taught by Reighard. Steere’s title changed to professor of systematic zoology, Reighard’s to professor of animal morphology. When Steere resigned in 1894, Reighard was appointed professor of zoology and director of the Zoological Laboratory and Museum. The Museum of Zoology (2015) 33

Reighard continued as head of the Department of Zoology until 1925. During this time, the biological sciences faculty at the University expanded and became more differentiated, in part due to the 1913 vote of the state legislature to fund construction of the Natural Science Building. In 1915, the Zoology Department moved into the new building and occupied approximately one-fourth of the space. A large number of new zoology faculty were recruited, spanning many emerging fields of research and teaching, including evolution, ecology, genetics, physiology, and development. The Museum also made three hires: Herbert Sargent (1898); Charles C. Adams (1903), who was recruited by Reighard to be curator of the Museum; and Alexander G. Ruthven (1906). These Museum positions initially lacked teaching duties, but Adams and Ruthven sought and obtained instructorships in the Department, and both individuals were instrumental in the development and flourishing of the UMMZ. C.C. Adams was a pioneering natural historian and field ecologist who initiated a series of extensive faunal and ecological surveys of the state that emphasized the importance of large sample sizes and detailed field data in addressing scientific questions. Ruthven was Adams’s Ph.D. student and became deeply interested in ecology and geographic distribution and adaptation, particularly of reptiles. In 1906, upon his graduation, Ruthven became Adams’s successor as curator of the University Museum, a position of leadership he held for 23 years.

1913-1956: Independence, Relocation, and Development

To Ruthven, museum collections and data were among the essential tools for understanding evolution and for constructing meaningful systematic relationships and taxonomies. A visionary scientist and administrator, Ruthven believed that museums should push in new directions to reflect changes in biological disciplines, and he was given that opportunity in 1913 when the disproportionate growth of the zoological collections prompted the regents to formally recognize the Museum of Zoology as a separate administration with Ruthven as director. From 1913-1956, the UMMZ reported to either the Board of 34 Museum of Zoology

Regents or the president of the University through its director or a museums operating committee. In addition to Ruthven, the full-time scientific staff of the newly recognized UMMZ included Norman A.Wood, as curator of birds; Crystal Thompson, scientific assistant in charge of fish and invertebrates; and Helen Thompson, scientific assistant in charge of amphibians and reptiles. Over the next eight years, Ruthven supervised the transformation of the UMMZ. The botany section was transferred to the newly formed University Herbarium, and a dynamic and far-reaching recruitment process reconstituted the UMMZ in six major divisions, each with at least one full-time curator: mammals (Lee Dice); birds (Wood); reptiles and amphibians (Ruthven, Helen Gaige); fishes (Carl Hubbs); insects (Frederick Gaige); and mollusks (Mina Winslow). It is notable that two of the UMMZ’s original seven curators were women. Ruthven was also highly adept at outreach, and under his leadership the Museum of Zoology attracted a large and active group of influential amateur naturalists. Informally known as the Detroit Naturalists Club, they participated in multiple aspects of the UMMZ program. Some members, such as Bryant Walker, E. B. Williamson, and William W. Newcomb, became nationally recognized authorities for specific taxonomic groups as well as honorary curators. A number were significant benefactors of the Museum of Zoology, supporting the establishment of the UMMZ publications series (Occasional Papers & Miscellaneous Publications), financing numerous national (especially regional) and international (especially neotropical) expeditions, and purchasing or donating important collections (e.g., B. Walker’s donation of his 100,000-lot private mollusk collection and 1,500-volume library). Indeed, a number of the UMMZ’s most important extant endowments (e.g., the Ammermann, Fargo and Swales funds) stem from the generosity of long-departed Detroit Naturalists Club members. The UMMZ’s collections and academic output were both rapidly expanding. The intensive investigation of Michigan’s fauna initiated under C.C. Adams continued with more emphasis on ecological relationships and biogeography. Targeted field expeditions were mounted all across the U.S. and The Museum of Zoology (2015) 35 into South America, Central America, and Mexico. By 1920, the UMMZ was growing out of the old Museum Building and overflowing into the Natural Science Building (for fishes, as Hubbs was an especially prolific collector), the Old Medical Building, and a succession of frame houses acquired by the University. In 1925, the legislature appropriated funds for a University Museums Building designed by the famous Detroit architect Albert Kahn—though Ruthven was centrally involved in the planning process. Finished in 1928, the new building was an innovative and exemplary early twentieth-century research university museum complex. It included space for the University Herbarium, the Museum of Anthropology, the Museum of Paleontology, a Section of Exhibits, and the Museum of Zoology. The new building brought administrative changes: Ruthven was appointed to the newly created post of director of University Museums and Frederick Gaige was made assistant director of the Museum of Zoology. Ruthven’s tenure was short- lived; in 1929 he was appointed president of the University. Gaige took his place and held the position until 1945. The Edwin S. George Reserve, situated just outside Ann Arbor, was provided as a gift to the University in 1930. It was placed under UMMZ administration and used as a venue for ecological research and teaching. Although Gaige’s directorship coincided with the Great Depression and World War II, the UMMZ’s collections and academic program continued to grow and develop during his tenure, with a high-quality training program that in many cases produced the next generation of curators both at the UMMZ and at peer museums across the country. Hubbs, in particular, proved to be a research and curatorial dynamo. He was interested in systematics, zoogeography, development, and the importance of hybridization in fish evolution, and by the time he left the UMMZ in 1944, the Museum of Zoology had become the leading ichthyological center in North America, training more than 40 Ph.D.s and growing the collection from 5,000 to almost 2.1 million specimens. (Hubbs’s student Reeve Bailey succeeded him in 1944.) Josselyn Van Tyne was appointed curator of birds in 1931 and 36 Museum of Zoology

initiated an era of tremendous growth in collections and in the ornithology program. Van Tyne and his students concentrated on life history and anatomical studies of birds, and Van Tyne acquired major private collections and established an outstanding ornithological library. In the mammal division, Lee Dice specialized in mammalian genetics, though in 1934 he became director of the Laboratory of Vertebrate Genetics, relinquishing his curatorship in 1938. William Burt, a behavioral ecologist, became curator of mammals in 1938 and was joined that year by assistant curator Emmett Hooper, a specialist in Mexican mammal diversity. Once Ruthven became University president, Helen Gaige assumed effective day-to-day responsibility for the Division of Herpetology. The UMMZ research and training program in reptiles and amphibians had become highly regarded and attracted many leading students, including some who would be recruited as divisional curators—including Norman Hartweg in 1934 and Charles Walker in 1947. Mina Winslow was curator of mollusks and engaged in regional and international collecting trips prior to her resignation in 1929. Winslow was replaced by one of the honorary mollusk curators, Calvin Goodrich, a specialist in North American pleurocerid snails who sampled extensively in southern U.S. drainages. In 1934, UMMZ graduate Henry van der Schalie was made assistant curator; he became curator upon Goodrich’s retirement in 1944. Van der Schalie established a prominent research and teaching program in snail-borne diseases and in mollusk biodiversity. Frederick Gaige’s research specialization concerned neotropical ants, but his research output was impacted by the demands of being director. The work of the insect division significantly expanded thanks to the activities of multiple honorary curators who took responsibility for major insect groups and added significantly to the collection. Two of these honorary curators, J. Speed Rodgers (Diptera) and T.H. Hubbell (Orthoptera), were UMMZ graduates who had obtained positions at the University of Florida. At the end of World War II, the UMMZ experienced an abrupt and significant loss of personnel. Frederick Gaige retired from the directorship and from his curatorship of insects and his spouse Helen Gage retired from her curatorship of herpetology. The Museum of Zoology (2015) 37

Their loss was keenly felt. The UMMZ turned to its former students to fill these and other gaps. J. Speed Rogers was appointed Museum director and T.H. Hubbell was appointed curator of insects. This pattern of hiring former UMMZ students continued during the ten years of Rodgers’ directorship; such appointments included Charles Walker as curator of herpetology in 1947 and Robert Miller as curator of fishes in 1948. Exceptions were the hiring in 1947 of William Gosline, an ichthyologist from Stanford University, and of Robert Storer in 1948, an ornithologist from University of California-Berkeley. The collections continued to grow significantly, as did the graduate training program.

1956-2011: No Longer an Island

Rodgers died unexpectedly in 1955 and was succeeded as director by T.H. Hubbell, who served in the role until his retirement in 1968. During his directorship, Hubbell reorganized and developed the UMMZ by broadening teaching and research programs; strengthened ties with the Zoology Department; made systematic biology more prominent in the curriculum; and added a research wing to the museum with monies from the National Science Foundation. The UMMZ and its sister museums had long faced teaching participation issues. Since 1928, teaching participation varied widely among the museums and even among individual curators, and all such participation was essentially ad hoc, voluntary, and associated with various departments (the Zoology Department in the case of the UMMZ). To regularize these practices, the UMMZ was made a department within the College of Literature, Science, and the Arts (LSA), reporting to the LSA dean and not to the University president or the Regents. UMMZ curators, in addition to retaining their research and curatorial functions in the Museum, expanded their roles in the undergraduate and graduate teaching program. Half of the academic-year appointments of the curators, teacher evaluations, and (most saliently) the tenure designations were transferred to the Department of Zoology. The UMMZ maintained partial independence by retaining half-time 38 Museum of Zoology

positions, and a separate budget and director. But the long-term fate and direction of the UMMZ was now inextricably tied to its new academic department. Hubbel appointed five new curators, including two future directors, who expanded the research questions and approaches the UMMZ would use, and renewed the teaching and training aspects of the program. Appointments included insect curator Thomas Moore (in 1956), a specialist in cicadas; and bird curator and former UMMZ student Harrison Tordoff in 1957. Tordoff studied Fringillidae systematics and crossbill genetics; he left in 1970 to join the Bell Museum at the University of Minnesota. Richard Alexander was recruited as curator of insects in 1957 and had a stellar career at the UMMZ, receiving numerous prestigious awards—including a Daniel Giraud Eliot Medal from the National Academy of Sciences in 1971—and being elected to that body in 1974. Alexander had broad interests in animal behavior and evolution. His early insect research focused on acoustic communication, speciation, and life history evolution. His later work addressed many central areas in evolutionary biology and attracted numerous graduate students, many of whom went on to successful scientific careers. In 1963, UMMZ graduate John B. Burch was appointed curator of mollusks. Burch developed an international research program on non-marine gastropods with an emphasis on species of medical importance and Pacific Island endemics as well as on karyological and immunotaxonomic characterization. Burch also founded a number of influential mollusk journals, most notably Malacologia, and currently serves as the UMMZ publications editor. Two new herpetology faculty were hired in the mid-1960s. Donald Tinkle, who was recruited from Texas Tech University in 1965 as a curator of amphibians and reptiles, was a pioneer in the use of long-term field experiments to study the life history of reptiles. In 1980, Tinkle was awarded the Eminent Ecologist Award of the Ecological Society of America, but he died that year from pancreatic cancer. Arnold Kluge was hired as a faculty member in the Department of Zoology in 1965—the year he was also given a research associate position in the UMMZ—and was upgraded to curator of amphibians and reptiles in 1967. The Museum of Zoology (2015) 39

Kluge’s interests included phylogenetic inference and, with Michigan botanist Herb Wagner and others, Kluge played an important role in the development of cladistics as a founding member of the Willi Hennig Society. In 1961, Hubbell applied to the National Science Foundation for major funding to construct a national facility for research in animal biosystematics. Thanks in part to the influential assistance of former curator Carl Hubbs, Hubbell was successful; this enabled construction of a new wing of UMMZ, opened in 1964. The UMMZ footprint significantly expanded, with a new lab, live animal facility, office, and a library that would quickly be fully occupied. In 1967, the UMMZ hosted a large international symposium on systematic biology sponsored by the National Research Council; the results were published in a 1969 volume by the National Academy of Sciences. The UMMZ was also active in international outreach. Most notably, Hubbell was instrumental in the 1963 formation of the inter-university Organization for Tropical Studies in Costa Rica with UMMZ curator Hartweg as its first president. Nelson Hairston served as UMMZ director from 1968 to1975. Appointed by LSA, he differed from his predecessors in that he was a faculty member in the Department of Zoology rather than a UMMZ curator. Hairston was a well-regarded ecologist who performed much of his fieldwork at UMMZ’s nearby Edwin S. George Reserve. Under his leadership, three new curators were recruited. Gerald Smith, a former UMMZ student, was appointed in 1969 as curator in both the UMMZ (Fishes) and in the Museum of Paleontology, and as assistant professor in the departments of Zoology and Geology. Smith worked with both extant and extinct fishes, especially in western drainages, and was particularly interested in rates of evolution and introgression. Before his retirement, he had the unique distinction of serving as director of three University Museums: Paleontology (1975-1981), Zoology (1998-2002), and the Herbarium (1999-2002). Robert Payne was appointed curator of birds in 1970, replacing Tordoff. Payne’s primary research interest was bird social behavior and systematics, with an emphasis on brood parasites and their hosts, especially African species. This 40 Museum of Zoology

involved both fieldwork in Africa and experimental behavior research with captive specimens at the UMMZ. Payne outlined a new rapid speciation mechanism caused by host switching associated with host song imprinting. He added large numbers of bird vocalization recordings as well as specimens to the UMMZ bird collection. Ronald Nussbaum was hired as curator of amphibians and reptiles in 1974, replacing Walker. He brought the number of herpetology curators to three. Nussbaum researched aspects of the evolution, ecology and systematics of amphibians and reptiles, especially taxa endemic to Africa, Seychelles, and Madagascar, and was extraordinarily active in documenting Malagasy reptile and amphibian diversity, including many unknown and threatened species. This work is of great conservation import and has added very significantly to the UMMZ herpetological collections. Donald Tinkle was UMMZ director from 1975 until his premature death in 1980 at the age of 50. His directorship coincided with an academic high water mark for the UMMZ. This was characterized by the production of a remarkable crop of graduate students under the advisorship of, among others, evolutionist Alexander, ecologist Tinkle, theorist Hamilton, bird behaviorist Payne, and cladist Kluge—and amplified by the presence of three or more curators per division. In October 1978, the Museum of Zoology celebrated its 50th year in the Ruthven Museums building by convening a symposium on natural selection and social behavior—areas to which curators and students had made important contributions. Some 700 individuals from all over the United States attended, and 31 formal papers were presented. The results were published in 1981 in a commemorative volume. Tinkle’s directorship also coincided with the 1975 merger of the Departments of Botany and of Zoology into a unified Division of Biological Sciences, a forerunner of the Department of Biology, with four divisions: botany; zoology; cell and molecular biology and ecology; and evolutionary and organismal Biology. William Dawson, a prominent animal physiologist, was appointed chair of the new department and served from 1975 to 1982. The Museum of Zoology (2015) 41

Two especially important hires were made during Tinkle’s five years as director of UMMZ. Philip Myers, initially a visiting curator of mammals in 1976, became an assistant curator the following year. Myers had a dual track career at the UMMZ. One concerned his research program on mammalian evolution and systematics, which focused on topics such as the evolutionary origins of Andean mammal diversity and on the long-term range dynamics of Michigan mammals associated with regional climate change. The other involved the origination and development of the highly innovative, award-winning Animal Diversity Web, a premier source of biodiversity information and science-learning exercises for K-12 and undergraduate students worldwide. In 1978, William Hamilton joined the UMMZ as curator of insects. Hamilton, an Englishman, was arguably the most prominent evolutionary theorist of his generation, famous for proposing a genetic basis for kin selection and altruism and winner of many prestigious awards—including, while at Michigan, foreign honorary membership in the American Academy of Arts and Science and fellowship in the Royal Society of London. The sociobiological implications of his theoretical work were controversial to some and led to student protests upon his arrival in Ann Arbor. In 1984, Hamilton was recruited back to the United Kingdom, where he accepted a position as Royal Society Research Professor at Oxford University. After Tinkle’s death, Robert Storer (curator of birds) served as interim director from 1980 to 1982. Two UMMZ curators were appointed during this period, including Barry OConnor as a curator of insects in 1980. Although he worked on insects, OConnor’s main research interest lay in Acari (mites) evolution, systematics, and taxonomy. Mites are extremely diverse, but relatively poorly studied. Many form commensal/parasitic associations with specific hosts and the vast majority of species remain undescribed. From a very modest base, over some 30 years, OConnor built one of the world’s great mite research collections in the UMMZ, obtaining NSF support to house his fast-growing microscope slide collection—a remarkable 42 Museum of Zoology

curatorial achievement. He also described many new species and helped to train a new generation of acarologists. William Fink was appointed curator of fishes in 1982, succeeding Reeve Bailey. His research on fish evolution and systematics combined a primary taxonomic focus on piranhas and other neotropical freshwater fishes, cladistic phylogenetics, and morphometric analyses of ontogenies. Fink was instrumental in organizing the 1988 Michigan Morphometric Workshop that resulted in a UMMZ special publication. He was an early adopter of museum catalogues computerization, obtaining multiple grants to create a database, place online, and network the UMMZ’s very large fish collection. Fink also trained the next generation of fish curators at many peer programs and served as UMMZ director from 2005 to 2011. In 1982, William Dawson completed his service as chair of the Division of Biological Sciences after seven years and was appointed director of the UMMZ; he served in that role until 1993. Molecular biology had developed to a point that it could be meaningfully applied outside of the discipline of biochemistry to address broader questions in the biological sciences. Dawson foresaw the value of incorporating a molecular perspective into UMMZ research and made a Laboratory of Molecular Systematics a prerequisite of his appointment. In 1980, Wesley Brown had joined the biology faculty as a visiting associate professor. When he was awarded tenure in1983, Dawson appointed him director of the UMMZ’s new Laboratory of Molecular Systematics. Brown was a pioneer in the use of mitochondrial DNA and protein sequence variation to address questions of broad evolutionary and phylogenetic significance. He attracted many ambitious postdoctoral researchers to his lab in the Natural Science Building and to the Laboratory of Molecular Systematics in the UMMZ, where they tackled a spectrum of high-profile evolutionary questions. The use of novel molecular approaches did not initially meet with unqualified enthusiasm across the broader UMMZ community, but within a decade molecular data was being routinely used in UMMZ research projects. In 1986, the Division of Biological Science was transformed into the Department of Biology, and The Museum of Zoology (2015) 43

Brown became chair of Biology from 1991 to 1996. As a result, Brown spent less time in the UMMZ and in the lab, instead relocating almost completely to the Natural Sciences Building. During Dawson’s directorship a number of tenure-track assistant curators did not obtain tenure, two in the mollusk division and one each in the bird and mammal divisions. All three vacant positions were eventually filled by candidates who had a significant molecular biology component to their research and who achieved tenure. One of the three, Priscilla Tucker, curator of mammals, was appointed by Dawson in 1988. Tucker is widely recognized for her evolutionary studies of Y chromosomes and for her detailed genetic deconstruction of a prominent European hybrid zone involving two closely related house mouse species that provide a model system for studying speciation in mammals. Tucker also significantly enhanced the UMMZ’s collection capabilities by obtaining NSF grant support to install the UMMZ’s first cryogenic facility, providing the capacity for high-quality preservation of specimen biomolecular structure. Dawson’s directorship also coincided with a period of significant NSF infrastructure grant support for collections in several UMMZ divisions, including fishes, mammals and reptiles, and amphibians. Richard Alexander assumed the UMMZ directorship from 1993 to 1998. He was deeply committed to the UMMZ and to the role of research museums in LSA academic programs. One of his first duties concerned searches for new bird and mollusk curators. This led to the appointment of David Mindell as curator of birds in 1994 and Diarmaid Ó Foighil as curator of mollusks in 1995. Both would become UMMZ directors. While at Michigan, Mindell had an active research program in bird molecular phylogeny, studying both ancient and more recent radiations, and he developed a parallel program in virus evolution. With Ó Foighil and others, Mindell purchased an ABI automated sequencer that gave UMMZ researchers access to cutting-edge sequencing capabilities. The sequencer was installed in the Laboratory of Molecular Systematics—renamed the Genomic Diversity Lab—with Mindell as its director. Mindell won NSF funding to upgrade the bird collection cabinets and subsequently served as UMMZ director for three 44 Museum of Zoology

years. He left Michigan in 2008 to become dean of the science and research collections at the California Academy of Sciences. Ó Foighil’s background was in marine mollusk evolution and systematics and his research continued in this vein, but he also branched out to work on freshwater and terrestrial taxa, with mollusk curator John Burch and his students. In particular, Ó Foighil used Burch’s samples of now-extirpated partulid tree snail populations to study the evolutionary history, systematics, and conservation biology of this endangered Pacific Island radiation. Ó Foighil obtained NSF funding to initiate computerization of the vast UMMZ mollusk collection. He became the director of UMMZ director and began a three-year term as EEB chair in 2014. Alexander’s UMMZ directorship unfolded against a background of growing discord within the Department of Biology, some of it aimed toward the associated museum units. This fractiousness was in some respects understandable. The Department had approximately 60 faculty spread across several buildings, making communication especially difficult between the two main blocks (essentially proto-departments with different cultures): ecology, evolution, and organismic biology, on the one hand, and molecular, cellular, and developmental biology on the other. Meanwhile, biology as a discipline was undergoing rapid expansion and differentiation with the emergence of many new specialized sub-fields. Departmental tensions were most apparent in prioritizing new hiring opportunities and the composition of search committees. Alexander believed that a certain constituency within the Department did not value Museum research and was communicating this perspective to the College. He feared that LSA would significantly cut the number of UMMZ curatorial lines and wrote a series of pointed reports to Dean Edie Goldenberg, outlining in considerable detail the value of University research museums in general and of the UMMZ’s program in particular. Gerald Smith served as director of UMMZ from 1998 to 2002 and as director of the Herbarium from an overlapping term of 2000 to 2002. This was a momentous period for the Department of Biology, the tenure home for UMMZ curators. The Museum of Zoology (2015) 45

Following an external review, LSA Dean Shirley Neuman in 2001 implemented a recommendation that the Department be split into two distinct entities. These became known as the Department of Molecular, Cellular, and Developmental Biology (MCDB) and the Department of Ecology and Evolutionary Biology (EEB). The two associated museum units migrated to EEB. This new department, led initially (and again later) by Deborah Goldberg, enjoyed a more cohesive academic environment and had a better collective appreciation for the importance of museum research. During Smith’s directorship, three UMMZ curators retired—Alexander and Moore, both of the insect division, and Burch, of the mollusk division. Smith worked assiduously with the College and Department to address these losses, leading to the hiring of new insect and mollusk curators, as well as the appointment of a manager for the Mollusk Collection—the only major UMMZ collection that lacked this support. Smith completed his service as director of UMMZ and the Herbarium in 2002. An external search was approved for a new director, but this proved unsuccessful. Curator David Mindell (birds) was appointed interim director for one year (2002-2003), then was renewed as director for a three-year term, of which only two were served (2003-2005). The two approved searches led to the appointment of L. Lacey Knowles as curator of insects in 2003 and Thomas Duda as curator of mollusks in 2004. Knowles was a global leader in the development of probabilistic methodologies for phylogenetic and phylogeographic analyses; her taxonomic focus was Orthoptera (grasshoppers and crickets). Knowles established a large, diverse, and highly productive research program at the UMMZ that attracted many students and post-docs and produced a steady stream of high-quality theoretical and empirical publications. Knowles and Laura Kobota hosted a NSF/UMMZ-funded Species Tree Workshop in January 2009 that attracted 170 participants interested in new approaches for estimating species trees. Knowles also co- edited the book that resulted, Estimating Species Trees: Practical and Theoretical Aspects, published in 2010. Duda is an innovative marine evolutionary biologist who has developed a highly 46 Museum of Zoology

integrative research program involving predatory Conus species: cone snails, one of the largest marine genera with hundreds of species. Duda studied the Conus radiation on many levels from a macroevolutionary perspective across the radiation to within- species analyses that link evolution at specific genes (paralyzing conotoxins) to organismal ecological performance (e.g., prey species repertoires). Duda also assembled a world-class collection of Conus species, including conchological, tissue, and expressed gene voucher material, as well as data on gut contents, genotype, and conotoxin repertoires. Smith (fishes) and Kluge (amphibians and reptiles) retired in 2003, leaving these two divisions relatively understaffed, with Fink and Nussbaum as sole curators for each. In 2004, Mindell submitted an ambitious five-year plan as director of both the UMMZ and the Herbarium. The plan proposed the unification of the UMMZ and Herbarium facilities with those of EEB in a single building or building complex on central campus. It also contemplated the establishment of an Institute for Systematics, Evolution and Biodiversity to help integrate the research, training, and public outreach activities of both museums and for the hiring of additional curators. The proposal did not meet with a receptive audience, in part because the College’s confidential long-term plans called for the relocation of research collections away from central campus. Following review of the five-year plans submitted by the Department and the museums, LSA decided to reduce the number of curators in the UMMZ from a total of 12 (in six FTEs) to eight (in four FTEs) and to formalize the reduction in the Herbarium that had already occurred from a peak of eight to four (in two FTEs). Mindell resigned from the UMMZ directorship in 2005. William Fink succeeded him from 2005 to 2011—a particularly eventful six-year period in the Museum’s history. Many aspects of the program were functioning well—individual research and teaching programs; Knowles’ successful Species Tree Workshop; networking of online vertebrate collection databases; establishment of the first cryogenic facility; and the continued success of the Animal Diversity Web. But attrition of the curatorial ranks continued with the retirement of bird curators The Museum of Zoology (2015) 47

Robert Payne and David Mindell in 2007 and 2008, respectively; this left one of the major UMMZ collections— with representatives of two-thirds of the planet’s bird species—without a curator or research and teaching program.EEB provided critical support in obtaining the College’s approval to conduct two searches during a period of very limited hiring opportunities—one in 2008 for a lower vertebrates organismal biologist, the other in 2009 for an evolutionary biologist. These held considerable promise for the UMMZ’s program. Unfortunately, neither search proved successful, although during the latter search a formal offer was extended to a highly qualified ornithologist couple. In 2009, a chronic and long-ignored safety issue regarding the UMMZ’s enormous ethanol-preserved collections, primarily fishes, reptiles and amphibians, finally came to a head. These world-class biodiversity collections were housed in facilities (some dating to the 1920s) that lacked modern fire suppression safety systems. Changing fire codes had put the ethanol- preserved collections in severe violation of new safety standards and the University decided to relocate them to a new facility that would meet current safety standards, adjacent to the Herbarium on Varsity Drive. This major initiative required a $17 million investment by the University in the UMMZ research collections, including the design and construction of new state- of-the-art ethanol collection facilities on Varsity Drive and (on a smaller scale) in the Ruthven Museums Building. The planning and construction phases alone required two years of detailed engagement and oversight by Fink, then the director, aided by the UMMZ’s team of collection managers. During 2009-2010, Fink was involved in a series of LSA- organized discussions with Deborah Goldberg, chair of EEB, and Paul Berry, director of the Herbarium, regarding inter- unit coordination. The College’s clear preference was that both museum units be merged with EEB, citing the need for better programmatic integration and for operational efficiencies. Following a formal joint external review of all three units in 2010, LSA proceeded with the merger, which was completed by July 2011. A detailed set of principles was agreed on, including recognition that curatorial care of the collections was a critical 48 Museum of Zoology

function of the Department and therefore curator responsibilities should continue to be associated with release from some teaching duties. Curators became 1.0 EEB faculty members, and the UMMZ’s four FTEs and Herbarium’s two FTEs were added to EEB’s total. But the title of curator was retained, and it was agreed that the number of faculty curators would be maintained at least at the level allocated by the College at the time of the merger (eight in the UMMZ, four in the Herbarium). The two museum director positions became EEB associate chairs, reporting to the EEB chair, but charged with responsibility for oversight of faculty curation, collection staff, and relevant operating budget and endowment funds.

2011-2013: A merger

Diarmaid Ó Foighil was appointed UMMZ director in July 2011. His most pressing task was to accomplish the complicated transfer of some five million biological specimens from the Ruthven Museums Building to the new Varsity Drive location. Construction was largely completed by December 2011, and the first specimens were moved to the new facility in January 2012. Thanks to a meticulously detailed planning process, the move itself went remarkably smoothly, with no specimens lost or damaged. This massive project was finished on August 3, 2012—under budget and seven weeks early, and the UMMZ’s impressive new ethanol-preserved facility has been fully operational since then. Transportation between the Ruthven and Varsity Drive location remains a logistical issue, but this has been ameliorated to some degree by an LSA-funded taxi service for UMMZ and Herbarium personnel, and by the use of LSA moving trucks for large consignments of ethanol-preserved specimens. When Daniel Rabosky became an EEB faculty member and UMMZ curator of amphibians and reptiles in late 2011-early 2012, it was an extremely positive development for the Department and the UMMZ biodiversity program. Rabosky was an emerging star in macroevolutionary studies whose research addressed fundamental aspects of among-lineage differences in species richness and phenotypic diversity, especially in The Museum of Zoology (2015) 49 vertebrates. His taxonomic focus was the Australian lizard and snake fauna. During his first year at the UMMZ, Rabosky won significant external funding, published high-profile papers, attracted students, taught innovative classes, and added significantly to the UMMZ reptile collection. Another important development for the EEB museum units was a significant upgrading of the electronic databases. Starting in 2011, the UMMZ and Herbarium directors participated in an LSA Collections Committee—initially led by Associate Dean Rick Francis—that reviewed practices for collection management, documentation, and digitization across all LSA museums. As a result, LSA committed to the purchase, installation, and support of the KE EMu collection software system for its museums. This represented a significant institutional commitment to the museums and the collections they curate, and marked a radical change from earlier practices in which each museum largely fended for itself. Perhaps the most significant development for EEB’s museum units in this period was LSA Dean Terrence McDonald’s announcement in fall 2011 of a “Transforming Ruthven” initiative. It was to relocate the collections and programs of the three Ruthven research museums (Anthropology, Paleontology, and Zoology) to Varsity Drive, joining the UMMZ ethanol- preserved collections and the Herbarium and thereby consolidating the University’s world-class natural history collections in one location. This initiative was coupled with plans to build a new biology building on central campus for MCDB and EEB faculty and their research programs, which was approved by the Regents in November 2014. In response to the dean’s announcement, the three Ruthven research museum directors and the Herbarium director wrote a memo outlining an integrated Varsity Drive plan that emphasized programmatic function needs and possible gains. For instance, with proper planning, the move to Varsity Drive would afford the opportunity to integrate collections more meaningfully into the undergraduate curriculum than had been possible in Ruthven to date. Many elements in that memo were incorporated into the Varsity Drive design. The UMMZ program was expected to move to its new Varsity 50 Museum of Zoology

Drive home by 2018. All collections databases were to migrate to KE Emu, the faculty curators were scheduled to be in a new Biology Building with their EEB colleagues for the first time; and the EEB program was to receive tens of millions of dollars in infrastructure support. EEB’s goal was to see its program emerge intact and functional—and among the best in the world.

Appendix: A Short History of the ESGR

The University of Michigan maintains the Edwin S. George Reserve (ESGR) for the purposes of providing research and education opportunities in the natural sciences and preserving its native flora and fauna. It is a 525-hectare fenced preserve located in Livingston County, Michigan, about twenty-five miles northwest of Ann Arbor. In 1927, to create a game preserve, George bought and fenced an area comprising eleven farms with tilled fields, small orchards, wood lots, pastures, and swamplands. In 1930, George donated the reserve to the University, which has maintained it since as a wildlife research area. The ESGR has been maintained by the University for the purposes of providing research and education opportunities in the natural sciences and preserving and demonstrating the native fauna and flora. It is used for natural history studies and other ecological investigations by Michigan students and staff and by visiting scientists from other institutions. From 1930 to 2005, the UMMZ was responsible for administering the ESGR. The last UMMZ curator to do so was Ronald Nussbaum, curator of amphibians and reptiles. There has been a rich history of long- and short-term biological research studies conducted on the ESGR. There also have been studies in other areas of the natural sciences, such as geology and atmospheric sciences. More than 425 research papers, 77 Ph.D. dissertations and 28 masters’ theses have been based on studies carried out wholly or in part in the ESGR. Long-term studies in the ESGR include an ongoing thirty-year study of turtle life histories and reproductive success that involves some records of individual turtles more than 70 years old; a long-term demographic study of the herd of white-tailed The Museum of Zoology (2015) 51 deer; a fifty-year study (1948-1997) study of successional changes in a 7.7-hectare abandoned field; and a 16-year study of the amphibian communities of 37 ponds. Shorter-term studies have addressed a wide range of ecological and evolutionary topics, including recent studies on bat echolocation; local and regional processes responsible for community patterns in dragonflies; paternity in white-tailed deer; trait-mediated indirect effects in food webs; melanism in peppered moths; intraguild predation in salamanders; the cost of slavery in ants; gypsy moth defoliation; and phenotypic plasticity in anuran larvae. In 2005 the administration of the ESGR was transferred to EEB;, EEB Professor Earl Werner served as the ESGR’s director from 2005 to 2012. He is a prominent ecologist who has studied community composition and dynamics in the reserve’s ponds for many years, collecting nearly 250,000 ecological specimens (including tadpoles, salamander larvae, fish, insects, mollusks, and crayfish) from thirty-eight ponds between 1996 and 2012. These specimens represent biodiversity vouchers for Werner’s influential body of published research on the ESGR pond ecosystems, and they have been formally placed in the UMMZ’s collections to ensure their availability for future generations of researchers. When Werner retired in 2013, the ESGR directorship passed to Christopher Dick, associate professor of EBB and Herbarium curator. — Earl Werner, edited by Diarmaid Ó Foighil, 2014