eis a ene, Mi a Perna, WaRORT OF Phe Beeman oF AEGIS OF a as Bh? FTG NOLAN i TP Bhd CPO 2 em BAG, BARR AE A Pricrpe errs tal foie busy Tea Renee gies Ae ks TTY SOR i ® Toate a my if ing I P nih wv a s 7 aah age a, v, : ; » - ; 5 7 ater Dek aro WOULD Haat in c y 7 : iw. ia at ; Nv 7 LY iw aan : aa) j a tytn i V , ra i) ; aD, r 7a | Ne ie Hrs 2 ey ee fi ch ANNUAL REPORT OF THE BOARD OF REGENTS OF THE SMITHSONIAN INSTITUTION SHOWING THE OPERATIONS, EXPENDITURES, AND CONDITION OF THE INSTITUTION FOR THE YEAR ENDED JUNE 30 L949 ’ (Publication 3996) UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON 1950 For sale by the Superintendent of Documents, U. S. Government Printing Office Washington 25, D. C. Price $2.75 (Buckram) a oT Bk Ook AR ASNGS Vin Ag, SMITHSONIAN INSTITUTION, Washington, December 9, 1949. To the Congress of the United States: In accordance with section 5593 of the Revised Statutes of the United States, I have the honor, in behalf of the Board of Regents, to submit to Congress the annual report of the operations, expendi- tures, and condition of the Smithsonian Institution for the year ended June 30, 1949. I have the honor to be, Respectfully, A. Wrermore, Secretary. BI SEP ~ 11950 a Sehee Lienntl CONTENTS Page Mii GeO bro LC ras tym nent Wings 4 Slvr els SE Aer hl en Sl cA ofS v Generalstatement see = eeunke so Se eet Bo 3 Be ee Lt) 1 Presentation of the Wright Brothers’ Aeroplane of 1903 to the United Statese Nationale Muse wit 2) 52 toe 2 oe eae eee ee ee Be ee 3 The establishment. ____..__-- ie kD Bay aioe ahd Leer ee aie alee ae SS a ao 5 Fiber BOasrdkotpecentcaa== pe tnt a5 as aie eee Bee a ee ey eee eee 6 PSSITY ATR COS eee ss AS Lh ay SR A 6 teen eoiiens = n a s ne e U IND PLOPMAt ONS = aes execs Soe Pee ae ee eee ee 7 WASILODS tet eps. aps BL win a hal ue Peres SH hen Re nt UO ET OE BA 8 Sixteenth James Arthur annual lecture on the sun___.___.___________-- 8 Summary of the year’s activities of the branches of the Institution_______ 9 Publications. = 222s 252s=22< fel 8a README eo AOE TY Li elt OL ICRRMS Shee Oe | SLES 14 ALS Altay see ol lan rs ein 1s Rs apres a ae OE aks 2 DAA septs SRE Lhd 15 Appendix 1. Report on the United States National Museum____________ 16 2. Report on the National Gallery of Arti! (22 20) 2S 2 25 3. Report on the National Collection of Fine Arts___________- 41 4 Reportontthesreem Gallery of Artes = sees ee ee eee 47 5. Report on the Bureau of American Ethnology____________- 55 6. Report on the International Exchange Service___________-_- 89 7. Report on the National Zoological Park_-_-______-______-- 97 8. Report on the Astrophysical Observatory ---____________-- 109 9: Report on-the National Air Museumis22-2 2222-25252 -_ === 114 10. Report on the Canal Zone Biological Area__--____________- 126 ileReport, onyiheslibrarys=22 == 720 shoe ea eee eae 132 25 Repongion pPublcabiong==2 263.993 2225s. - ose ode eee 136 Report of the executive committee of the Board of Regents________-____- 143 GENERAL APPENDIX hestormavionsolestarsbyaliyInanyoDILZCl dae ae eee 153 Aloe Cream OI ToS Capel, loyy ANoveriao leRVE ee eee enneeeoe 161 The 200-inch Hale telescope and some problems it may solve, by Edwin Pole eas ee eee eye oe ee ee ee eye Ae ake 3 175 The determination of precise time, by Sir Harold Spencer Jones________- 189 The elementary particles of physics, by Carl D. Anderson_____-_____--- 203 Recent advances in virus research, by Wendell M. Stanley___-__-___--_-- 213 Ground-water investigations in the United States, by A. N. Sayre____--- 219 Modern soil science, by Charles E. Kellogg_.____._._-...---------------- 227 Aime in-ovolution bys EeZeUnere cejsee= oss se- Sao ao cal ee cess 247 More about animal behavior, by Ernest P. Walker___-__________------- 261 IV CONTENTS Page The breeding habits of the weaverbirds: a study in the biology of behavior patterns, by dlerbert Priedmann--..= 222252522 -s2 oo See eee ee 293 New Zealand, a botanist’s paradise, by Egbert H. Walker______________ 317 The archeological importance of Guatemala, by A. V. Kidder__________- 349 Excavations at the prehistoric rock-shelter of La Colombiére, by Hallam MGSO WIS Bisa 5, olan eo Ds ee ee ee ee pee 359 Ronne Antarctic research expedition, 1946-1948, by Commander Finn Ronne aU Ore Neg hes. oe a te 2 Se ee ee ee ee pe ee 369 whe staterot science, by Karl. ©, Compton=c- 22- = ee eee 395 LIST OF PLATES Secretary 6. report: Lu atOs two: ess sk ee 2s Se a eee ee 54 ihe formation of stars (Spitzer blates) jae 2) fe see ee ee one ee 160 Whe origin of theyearth.@Page):, Plates Iai! ss. 2ei eee Se 166 The 200-inch Hale telescope (Hubble): Plates 1-10________________-_-- 182 Modern soil science (Kellogg): Plates 1,2. 225-5. 8 ee 246 Animal behavior (Ernest Walker): Plates 1-16._..._..........._.._--- 278 The breeding habits of the weaverbirds (Friedmann): Plates 1-8_______- 310 New Zealand, a botanist’s paradise (Egbert Walker): Plates 1-10______- 326 The archeological importance of Guatemala (Kidder): Plates 1-6__-_____- 358 Excavations at La Colombiére (Movius): Plates 1-7_____.._--_________- 366 Ronne Antarctic research expedition (Ronne): Plates 1-8_----_____--_--- 374 THE SMITHSONIAN INSTITUTION June 30, 1949 Presiding Officer ex officto— Harry S. Truman, President of the United States. Chancellor.—Frep M. Vinson, Chief Justice of the United States. Members of the Institution: Harry 8S. Truman, President of the United States. ALBEN W. BaRKLEY, Vice President of the United States. Frep M. Vinson, Chief Justice of the United States. Grorecs C. MARSHALL, Secretary of State. Joun W. Snyper, Secretary of the Treasury. Louis JoHNSON, Secretary of Defense. Tom C. Cuarx, Attorney General. JessE M. Donatpson, Postmaster General. Jutius A. Krua, Secretary of the Interior. CHARLES F, BRANNON, Secretary of Agriculture. CHARLES SAWYER, Secretary of Commerce. Maurice Tosin, Secretary of Labor. Regents of the Institution: Frep M. Vinson, Chief Justice of the United States, Chancellor. ALBEN W. BARKLEY, Vice President of the United States. Cuinton P. ANDERSON, Member of the Senate. LEVERETT SALTONSTALL, Member of the Senate. Wat7TEerR F. Grorce, Member of the Senate. CLARENCE Cannon, Member of the House of Representatives. E. E. Cox, Member of the House of Representatives. Joun M. Vorys, Member of the House of Representatives. Harvey N. Davis, citizen of New Jersey. Arraur H. Compton, citizen of Missouri. VANNEVAR Busu, citizen of Washington, D. C. Rosert V. Fueming, citizen of Washington, D. C. JEROME C. HuNSsAKER, citizen of Massachusetts. Executive Commitiee—Ropert V. FLEMING, chairman, VANNEVAR CLARENCE CANNON. Secretary — ALEXANDER WETMORE, Assistant Secretary. JOHN E. Grar. Assistant Secretary.—J. L. Keppy. Administrative assistant to the Secretary.—LouisE M, PEARSON. Treasurer.—J. D. Howarp. Chief, editorial division. WeEBsTER P. TRUE. Librarian.—Leiua F, Cuarx. Administrative accountant.—TuHomas F, Cuark. Superintendent of buildings and labor.—L. L. OLIVER. Personnel officer —B. T. CARWITHEN. Chief, division of publications.—L. E. COMMERFORD. Property, supply, and purchasing officer—ANTHONY W, WILDING. Photographer.—F. B. Kustner. Busi, VI ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 UNITED STATES NATIONAL MUSEUM Director.—A. REMINGTON KELLOGG. Chief, office of correspondence and records—HrLeNA M. WEIss. Editor.—Pauu H. OEHSER. Assistant librarian.— ELIsABETH H. Gazin. SCIENTIFIC STAFF DEPARTMENT OF ANTHROPOLOGY: Frank M. Setzler, head curator; A. J. Andrews, chief preparator. Collaborator in anthropology: W. W. Taylor, Jr. Division of Archeology: Neil M. Judd, curator; Waldo R. Wedel, associate curator; M. C. Blaker, scientific aid; J. Townsend Russell, honorary assistant curator of Old World archeology. Division of Ethnology: H. W. Krieger, curator; J. C. Ewers, associate curator; C. M. Watkins, associate curator; R. A. Elder, Jr., assistant curator. Division of Physical Anthropology: T. Dale Stewart, curator; M. T. Newman, associate curator. DEPARTMENT OF ZOOLOGY: Waldo L. Schmitt, head curator; W. L. Brown, chief taxidermist; Aime M. Awl, scientific illustrator. Associates in Zoology; T. 8S. Palmer, W. B. Marshall, A. G. Béving, C. R. Shoemaker, W. K. Fisher. Collaborator in Zoology: R. 8. Clark. Collaborator in Biology: D. C. Graham. Division of Mammals: D. H. Johnson, associate curator; H. W. Setzer, asso- ciate curator; N. M. Miller, museum aid; A. Brazier Howell, collaborator; Gerrit S. Miller, Jr., associate. Division of Birds: Herbert Friedmann, curator; H. G. Deignan, associate curator; Alexander Wetmore, custodian of alcoholic and skeleton collec- tions; Arthur C. Bent, collaborator. Division of Reptiles and Amphibians: Doris M. Cochran, associate curator. Division of Fishes: Leonard P. Schultz, curator; E. A. Lachner, associate curator; L. P. Woods, associate curator; D. 8. Erdman, scientific aid; W. T. Leapley, museum aid. Division of Insects: L. O. Howard, honorary curator; Edward A. Chapin, curator; R. E. Blackwelder, associate curator; W. D. Field, associate curator; O. L. Cartwright, associate curator; Grace E. Glance, associate curator; W. L. Jellison, collaborator. Section of Hymenoptera: S. A. Rohwer, custodian; W. M. Mann, assist- ant custodian; Robert A. Cushman, assistant custodian. Section of Diptera: Charles T. Greene, assistant custodian. Section of Coleoptera: L. L. Buchanan, specialist for Casey collection. Section of Lepidoptera: J. 'T. Barnes, collaborator. Division of Marine Invertebrates: F. A. Chace, Jr., curator; P. L. Illg, asso- ciate curator; Frederick M. Bayer, assistant curator; L. W. Peterson, G. S. Cain, museum aids; Mrs. Harriet Richardson Searle, collaborator; Max M. Ellis, collaborator; J. Percy Moore, collaborator; Mrs. M. 8. Wilson, collaborator in copepod Crustacea. Division of Mollusks: Harald A. Rehder, curator; Joseph P. E. Morrison, associate curator; R. Tucker Abbott, assistant curator; W. J. Byas, museum aid; P. Bartsch, associate. Section of Helminthological Collections: Benjamin Schwartz, collabo- rator, Division of Echinoderms: Austin H. Clark, curator. SECRETARY’S REPORT VII DEPARTMENT OF Borany (Nationant HERBARIUM): EK. P. Killip, head curator; Henri Pittier, associate in botany. Division of Phanerogams: A. C. Smith, curator; E. C. Leonard, associate curator; E. H. Walker, associate curator; Lyman B. Smith, associate curator; V. E. Rudd, assistant curator. Division of Ferns: C. V. Morton, curator. Division of Grasses: Jason R. Swallen, curator; Agnes Chase, research asso- ciate; F. A. McClure, research associate. Division of Cryptogams: E. P. Killip, acting curator; Paul S. Conger, asso- ciate curator; G. A. Llano, associate curator; John A. Stevenson, custodian of C. G. Lloyd mycological collections; W. T. Swingle, custodian of Higher Algae; David Fairchild, custodian of Lower Fungi. DEPARTMENT OF GEOLOGY: W. F. Foshag, head curator; J. H. Benn, exhibits preparator; Jessie G. Beach, aid. Division of Mineralogy and Petrology: W. F. Foshag, acting curator; E. P. Henderson, associate curator; G. S. Switzer, associate curator; F. E. Holden, exhibits preparator; Frank L. Hess, custodian of rare metals and rare earths. Division of Invertebrate Paleontology and Paleobotany: Gustav A. Cooper, curator; A. R. Loeblich, Jr., associate curator; David Nicol, associate curator; W. T. Allen, L. Pendleton, museum aids; J. Brookes Knight, research associate in Paleontology. Section of Invertebrate Paleontology: T. W. Stanton, custodian of Mesozoic collection; J. B. Reeside, Jr., custodian of Mesozoic collection. Division of Vertebrate Paleontology: C. L. Gazin, curator; D. H. Dunkle, asso- ciate curator; Norman H. Boss, chief exhibits preparator; W. D. Crockett, scientific illustrator; A. C. Murray, F. L. Pearce, preparators. Associates in Mineralogy: W. T. Schaller, S. H. Perry, J. P. Marble. Associates in Paleontology: T. W. Vaughan, R. S. Bassler. DEPARTMENT OF ENGINEERING AND INDUSTRIES: Frank A. Taylor, head curator. Division of Engineering: Frank A. Taylor, acting curator. Section of Civil and Mechanical Engineering: Frank A. Taylor, in charge. Section of Marine Transportation: Frank A. Taylor, in charge. Section of Electricity: K. M. Perry, associate curator. Section of Physical Sciencesand Measurement: Frank A. Taylor, in charge. Section of Land Transportation: S. H. Oliver, associate curator. Division of Crafts and Industries: W. N. Watkins, curator; F. C. Reed, associate curator; E. A. Avery, museum aid; F. L. Lewton, research associate. Section of Textiles: G. L. Rogers, assistant curator. Section of Wood Technology: William N. Watkins, in charge. Section of Manufactures: F. C. Reed, in charge. Section of Agricultural Industries: F. C. Reed, in charge. Division of Medicine and Public Health: G. S. Thomas, associate curator. Division of Graphic Arts: J. Kainen, curator; E. J. Fite, museum aid. Section of Photography: A. J. Wedderburn, Jr., associate curator. DEPARTMENT OF History: Charles Carey, acting head curator; T. T. Belote, Museum historian. Divisions of Military History and Naval History: M. L. Peterson, associate curator; J. R. Sirlouis, assistant curator. Division of Civil History: M. W. Brown, assistant curator. Division of Numismatics: 8S. M. Mosher, associate curator. Division of Philately: C. L. Manning, assistant curator. VIII ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 NATIONAL GALLERY OF ART Trustees: Frep M. Vinson, Chief Justice of the United States, Chairman. Grorcse C. MarsHatt, Secretary of State. Joun W. Snyper, Secretary of the Treasury. ALEXANDER WETMORE, Secretary of the Smithsonian Institution. SaMvuEL H. Kress. FERDINAND LAMMOT BELIN. DuNCAN PHILLIPS. CHESTER DALE. Paut MELLon. President.—SAMUEL H. Kress. Vice President—FERDINAND LAMMOT BELIN. Secretary-Treasurer.—HUNTINGTON CAIRNS. Director.—Davip E. FINLEY. Administrator.—Harry A. McBrive. General Counsel. HUNTINGTON CAIRNS. Chief Curator—JoHN WALKER. Assistant Director.—MacGILL JAMES. NATIONAL COLLECTION OF FINE ARTS Director.—Tuomas M. Braces. Curator of ceramics.—P. V. GARDNER. Exhibits preparator.—G. J. Martin. Assistant librarian.— ANNA M. LINK. FREER GALLERY OF ART Director.—A. G. WENLEY. Assistant Director.—JoHN A. PoPpE. Associate in Near Eastern art.—RicHARD ETTINGHAUSEN. Associate in Far Eastern art.—W. R. B. AcKER. Research associate—Gracr DUNHAM GUEST. BUREAU OF AMERICAN ETHNOLOGY Director.—MatTTHEWw W. STIRLING. Associate Director —F RANK H. H. Roserts, Jr. Senior ethnologists—H. B. Couuins, Jr., Joan P. Harrineton, W. N. FENTON. Senior anthropologists —G. R. Wituny, P. Drucker. Collaborators —FRANcES Densmore, JoHn R. Swanton, A. J. WaRina, Jr. Editor—M. HELEN PALMER. Assistant librarian.—Miriam B. KetcHum. Scientific illustrators —Epwin G. Cassnpy, E. G. SCHUMACHER. Archives assistant.—M. W. Tucker. INsTITU1E OF SocraL ANTHROPOLOGY.—G. M. Fostrmr, Jr., Director. River Basin SurvEyS.—F RANK H. H. Rosmrts, Jr., Director. INTERNATIONAL EXCHANGE SERVICE Chief.—D. G. WILLIAMs. SECRETARY’S REPORT Ix NATIONAL ZOOLOGICAL PARK Director.—Wiiu1aAM M. Mann. Assistant Director.—Ernest P. WALKER. Head Keeper.—F Rank O. Lowe. ASTROPHYSICAL OBSERVATORY Director.—Loyat B. ALDRICH. Assistant librarian.— Marjorie KuNzE. Division oF ASTROPHYSICAL RESEARCH: Chief.—Wi.ii1amM H. Hoover. Instrument makers.—ANDREW KRAMER, D. G. Tatsert, J. H. Harrison, Research associate-—CHARLES G, ABBOT. DIvIsION OF RADIATION AND ORGANISMS: ~ Chief —R. B. WirHrow. Plant physiologist (physicochemical). —LrONARD PRICE. Biological aid (botany).—Y. B. Ensrap. NATIONAL AIR MUSEUM Advisory Board: ALEXANDER WETMORE, Chairman. Mas. Gen. GRANDISON GARDNER, U.S. Air Force. Rear Ap. A. M. Prinz, U. S. Navy. Grover LOENING. WixuiaM B. Srovut. Assistant to the Secretary for the National Air Musewm.—Caru W. Mirman, Curator.—P. E. GARBER. Associate curators.—S. L. Brrers, R. C. Srrospet, W. M. Mates. Exhibits preparator.—S. L. Porter. CANAL ZONE BIOLOGICAL AREA Resident Manager.—Jamus ZETEK. ty o Lien 7 ow REPORT OF THE SECRETARY OF THE SMITH- SONIAN INSTITUTION ALEXANDER WETMORE FOR THE YEAR ENDED JUNE 30, 1949 To the Board of Regents of the Smithsonian Institution: GENTLEMEN: I have the honor to submit herewith my report show- ing the activities and condition of the Smithsonian Institution and its bureaus during the fiscal year ended June 30, 1949. GENERAL STATEMENT The Institution continued vigorously to pursue its program of activities in ‘‘the increase and diffusion of knowledge” as stipulated by its founder, James Smithson. The increase of knowledge is fos- tered by original scientific researches and explorations in the fields of anthropology, biology, geology, and astrophysics; the diffusion of knowledge, by publications in a number of series that are distributed free to libraries and educational institutions throughout the world, by extensive museum and art gallery exhibits, by the International Exchange Service for the world-wide interchange of scientific and governmental publications, and by a large correspondence, both national and international. I present first certain general features of the year’s activities, together with a summary of the work of the several bureaus of the Institution, to afford a concise picture of the events of the year. Next follow appendixes containing more detailed reports on each bureau, and finally there appears the financial statement of the Execu- tive Committee of the Board of Regents. The appendixes contain reports on the United States National Museum, the National Gallery of Art, the National Collection of Fine Arts, the Freer Gallery of Art, the Bureau of American Ethnology, the International Exchange Service, the National Zoological Park, the Astrophysical Observatory, the National Air Museum, the Canai Zone Biological Area, the Smith- sonian library, and the publications of the Institution. 1 2 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 When the Smithsonian Institution began its operations more than one hundred years ago, it carried on its research programs largely by subsidizing the work of scientists not on its own staff, and by publish- ing the results of their work. As these pioneer researches expanded in scope and became somewhat stabilized, bureaus gradually grew up around the Institution, each with its own staff specializing in the work of that particular field. The value of the various activities gradually became known to the Nation, and eventually one by one they were recognized as public necessities by the Congress. Most of them are now supported largely by Government funds although re- maining under Smithsonian direction. At present, nearly all the research and exploration of the Institution is done through these bureaus, notably the United States National Museum, the Bureau of American Ethnology, and the Astrophysical Observatory. As stated in last year’s report, the Institution has for many years operated under the handicap of shortages of personnel and of ade- quate housing space. I reported that the Smithsonian Institution has today the same amount of space that it had in 1911 in which to accommodate four times as many visitors and four times as many museum specimens. Much the same condition still prevails. Some slight gain was apparent in personnel in a few of the scientific divi- sions, but not sufficient for the prompt execution of essential cura- torial work and adequate research on the National collections. The crowded condition, particularly in the buildings of the National Museum, remained unalleviated. In the report of the Director of the Museum it will be noted that there is a considerable decrease in number of specimens accessioned during the year, a decrease which, he says, “may be attributed in part to the inadequacy of available storage facilities for the preservation of such materials.”’ More ade- quate building space is one of our major needs. Though hampered by space conditions it should be brought to attention that the Smithsonian Institution continues to grow and to expand its usefulness year by year. In the 5 years during which I have served as Secretary, three additional activities have been added to its responsibilities—the Canal Zone Biological Area, the National Air Museum, and the River Basin Surveys, the latter a unit of the Bureau of American Ethnology. The work of these new activities has notably augmented Smithsonian efforts toward the increase and diffusion of knowledge in widely diversified fields, as will be seen in reading the detailed reports appended hereto. The purpose in calling attention to deficiencies is to emphasize the obvious fact that a growing SECRETARY’S REPORT 3 institution such as the Smithsonian, of so vital interest and importance to the American people, must receive increased financial support if it is to continue to meet its full obligations and to further the high ideals of its founder, James Smithson, who left his entire fortune in trust to the United States of America for the benefit of all mankind; PRESENTATION OF THE WRIGHT BROTHERS’ AEROPLANE OF 1903 TO THE UNITED STATES NATIONAL MUSEUM On December 17, 1948, the forty-fifth anniversary of the first flight by Wilbur and Orville Wright at Kitty Hawk, N. C., the original aeroplane that made that historic flight became the property of the American people. At a formal ceremony in the Museum attended by many high civil and military officials the plane was presented to the United States National Museum by Milton Wright on behalf of the estate of Orville Wright. The story of the plane goes back to December 17, 1903, when the Wright Brothers were ready after several years of research and experiment to test out their gasoline-engine-powered biplane at Kitty Hawk on the coast of North Carolina. With Orville at the controls, the machine was released, and after a 40-foot run on the launching track, it lifted into the air in full flight. In Orville Wright’s own words: “The flight lasted only 12 seconds, but it was nevertheless the first in the history of the world in which a machine carrying a man had raised itself by its own power into the air in full flight, had sailed forward without reduction of speed, and had finally landed at a point as high as that from which it started.” Three more flights were made the same day, but after the last flight a strong gust of wind turned the plane over, damaging it so badly that no more trials were made that year. The damaged machine and engine were sent back to the Wrights’ workshop in Dayton, and 13 years later were restored, using all the original parts available. The aeroplane was displayed at the Massachusetts Institute of Technology and later at several aeronautical exhibitions. In 1928 Orville Wright had it sent as a loan to the Science Museum at South Kensington, London, England, where it remained on exhibi- tion until World War II. Owing to the danger of damage by bomb- ing, the plane was removed to a safe place for the duration of the war, When Orville Wright died on January 30, 1948, it was learned from papers in his files that he wished the Kitty Hawk aeroplane to be returned to the United States and placed in the National Museum. The executors of his estate conferred with officials of the Science 4 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Museum and of the Smithsonian Institution, and with the generous cooperation of the British Government the actual transfer of the plane took place in November 1948. It was brought across the Atlantic to Halifax on the Mauretania, from there to Bayonne, N.J., on the Navy carrier Palau, and to Washington by Navy truck. At the formal presentation on December 17, 1948, the ceremonies were opened by the Secretary of the Smithsonian Institution. After the invocation by Maj. Gen. Luther D. Miller, Chief of Chaplains, Department of the Army, and greetings by the Presiding Officer, Chief Justice Fred M. Vinson, Chancellor of the Smithsonian Institu- tion, a message from the President of the United States was read by Col. Robert B. Landry, Air Force Aide to the President. His Britan- nic Majesty’s Ambassador, Sir Oliver Franks, K. C. B., C. B. E., then spoke on “Britain and the Wright Brothers,” after which the presentation of the aeroplane was made by Milton Wright, of Dayton, Ohio, on behalf of the estate of Orville Wright. Mr. Wright told of his boyhood recollections of his uncles’ bicycle shop where the Kitty Hawk plane was fabricated, and concluded thus: “The aeroplane means many things to many people. To some it may be a vehicle for romantic adventure or simply quick transporta- tion. To others it may be a military weapon or a means of relieving suffering. To me it represents the fabric, the glue, the spruce, the sheet metal, and the wire which, put together under commonplace circumstances but with knowledge and skill, gave substance to dreams and fulfillment to hopes.” The aeroplane was accepted on behalf of the Smithsonian Institu- tion by Chief Justice Fred M. Vinson, Chancellor of the Institution, and the address of acceptance was given by Vice President-Elect Alben W. Barkley, a regent of the Institution. In the course of his address Mr. Barkley expressed one thought that doubtless was in the minds of all participants in the ceremony: “Tt is a matter of deep regret to all of us that Orville Wright could not have been here today to see this wide public recognition of achieve- ment, and receive in person the fitting acclaim to his brother, to himself, and to their Kitty Hawk plane. We are grateful to all of those who have made it possible to bring the plane back to its native soil, and especially to the heirs of the estate of Orville Wright, for depositing the Kitty Hawk machine here where all America will have an opportunity to see it, and where all may do it fitting honor.” The Kitty Hawk aeroplane now hangs suspended from the ceiling of the north hall of the National Museum’s Aris and Industries Building, where the presentation ceremony was held. Directly back of the main entrance, the plane is the first object to meet the eyes of SECRETARY’S REPORT 5 the thousands of visitors who throng the Museum daily. As thus displayed it bears the following label: The Original WRIGHT BROTHERS’ AEROPLANE The world’s first power-driven heavier-than-air machine in which man made free, controlled, and sustained flight Invented and built by Wilbur and Orville Wright Flown by them at Kitty Hawk, North Carolina December 17, 1903 By original scientific research the Wright Brothers discovered the principles of human flight As inventors, builders, and flyers they further developed the aeroplane, taught man to fly and opened the era of aviation Deposited by the Estate of Orville Wright ° “The first flight lasted only twelve seconds, a flight very modest compared with that of birds, but it was nevertheless the first in the history of the world in which a machine carrying a man had raised itself by its own power into the air in free flight, had sailed forward on a level course without reduction of speed, and had finally landed without being wrecked. The second and third flights were a little longer, and the fourth lasted 59 seconds covering a distance of 852 feet over the ground against a 20 mile wind.’”—W1.LzBur and OrvILLE WriGcaHt. (From Century Magazine, vol. 76, September 1908, p. 649.) This is not the final resting place of the plane, however—it is destined eventually to occupy the place of honer in the National Air Museum, the most recent bureau of the Smithsonian Institution. Preliminary plans for the Air Museum envision a special centrally located exhibit area for the Wright aeroplane of 1903, to serve as a memorial to the birth of aviation. THE ESTABLISHMENT The Smithsonian Institution was created by act of Congress in 1846, according to the terms of the will of James Smithson, of England, who in 1826 bequeathed his property to the United States of America “to found at Washington, under the name of the Smithsonian Insti- tution, an establishment for the increase and diffusion of knowledge among men.’ In receiving the property and accepting the trust, Congress determined that the Federal Government was without authority to administer the trust directly, and, therefore, constituted an ‘establishment’? whose statutory members are “the President, the Vice President, the Chief Justice, and the heads of the executive departments.” 6 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 THE BOARD OF REGENTS The following changes occurred during the year in the personnel of the Board of Regents: On January 20, 1949, Vice President Alben W. Barkley (formerly a regent by appointment from the Senate) became ‘ex officio a member of the Board. On February 14, 1949, the following regents were appointed from the House of Representatives: Clarence Cannon of Missouri; John M. Vorys of Ohio; and E. E. Cox of Georgia to succeed Samuel K. McConnell of Pennsylvania. On March 8, 1949, Senators Leverett Saltonstall and Clinton P. Anderson were appointed to succeed Vice President Alben W. Barkley who became an ex officio member of the Board, and Senator Wallace H. White of Maine, retired. On March 10, 1949, Dr. Jerome C. Hunsaker was appointed a citizen regent from Massachusetts for the statutory term of 6 years, to succeed Frederic C. Walcott, retired. The roll of regents at the close of the fiscal year, June 30, 1949, was as follows: Chief Justice Fred M. Vinson, Chancellor; Vice President Alben W. Barkley; members from the Senate: Walter F. George, Clinton P. Anderson, Leverett Saltonstall; members from the House of Repre- sentatives: Clarence Cannon, John M. Vorys, E. E. Cox; citizen mem- bers: Harvey N. Davis, Arthur H. Compton, Vannevar Bush, Robert V. Fleming, and Jerome C. Hunsaker. Proceedings.—The Board of Regents held its annual meeting on January 14,1949. Present: Chief Justice Fred M. Vinson, Chancellor; Representative Clarence Cannon, Representative John M. Vorys, Dr. Arthur H. Compton, Dr. Harvey N. Davis, Dr. Robert V. Flem- ing, Secretary Alexander Wetmore, and Assistant Secretary John E. Graf. The Secretary presented his annual report covering the activities of the Institution and its bureaus, including the financial report of the Executive Committee, for the fiscal year ended June 30, 1948, which was accepted by the Board. The usual resolution authorizing the expenditure by the Secretary of the income of the Institution for the fiscal year ending June 30, 1950, was adopted by the Board. It was announced that in support of the work of the Astrophysical Observatory John A. Roebling had made a further generous gift which is of major importance in carrying on these scientific investi- gations. The annual report of the Smithonsian Art Commission was pre- sented by the Secretary and accepted by the Board. A resolution was adopted to reelect the following members for 4-year terms: Archibald SECRETARY’S REPORT he G. Wenley, David E. Finley, Eugene E. Speicher, Paul Manship. The following officers were reelected for the ensuing year: Chairman, Paul Manship; vice chairman, Robert Woods Bliss; secretary, Alexander Wetmore. The Board was advised that in an attempt to recover the Gellatly art collection from the Secretary in his status of a private individual, though acting as custodian under the Smithsonian Institution, Mrs. Charlayne Gellatly’s attorneys had filed action in the District Court of the United States for the District of Columbia on June 18, 1947. Under date of June 17, 1948, Judge J. McGuire rendered a decision that, in the opinion of the Court, there was no merit in Mrs. Gellatly’s claims, since it was found that there was a valid gift to the United States by the deceased, John Gellatly, before his death and before his marriage. On July 19, 1948, the attorney for Mrs. Gellatly filed notice of appeal before the United States Court of Appeals for the District of Columbia. Marvin C. Taylor, special attorney, Department of Justice, represented the Institution. On the evening of March 1, 1949, an informal meeting of the Board was held at dinner in the Main Hall of the Smithsonian Building, with the Chancellor, Chief Justice Fred M. Vinson, presiding. At this meeting heads of the various activities under the Institution pre- sented statements relative to their work. These statements, with the ensuing discussion, provided a general view of the existing operations of the Smithsonian, particularly in the research field. FINANCES A statement on finances, dealing particularly with Smithsonian private funds, will be found in the report of the Executive Committee of the Board of Regents, page 143. APPROPRIATIONS Funds appropriated to the Institution for the fiscal year ended June 30, 1949, totaled $2,259,000, allotted as follows: @eneraleadministra tions sae see ee ee ee ee $46, 794 INationale@iuse ini fae ee ae eee ee ee eg 712, 560 Buresawvof American hthuolopy ss 222 22225 2 eee ee 71, 996 AstrophysicalObsernvatorye. sesso etek ot sete - caer 101, 590 INationalaCollectionsoteliner Arts == eee ee ee ee eee 32, 543 International Exchange Service=* 22 v. oul sas See be ee ole 68, 938 Maintenance and-OperaviOn. 2.052) nee eee eo eo 764, 626 Dervicerdivisions=seses see ee ee ee ee 274, 448 INationaleAirsMiniscn mee ee ei ee ee ee ee eee 180, 285 CanalkZoneybiolocicalwArca to 3 ins so = ee oe ee eee 4, 760 Wnallottedses: -tee See Seek! Wha BEM NE esa A ee ae a Oe 460 DLT C come 2 a ce ld Se pA ee rt eA Ne PE TR ea 2, 259, 000 866591—50——-2 8 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 In addition $1,073,500 was appropriated to the National Gallery of Art, a bureau of the Institution but administered by a separate board of trustees; and $528,848 was provided in the District of Columbia appropriation act for the operation of the National Zoological Park. Besides these direct appropriations, the Institution received funds by transfer from other Federal agencies, as follows: From the State Department, from the appropriation Cooperation with the American Republics, 1949, a total of $97,900 for the operation of the Institute of Social Anthropology, including the issuance of publications resulting from its work. From the National Park Service, Interior Department, $118,500 for archeological projects in connection with River Basin Surveys. VISITORS The number of visitors to the Smithsonian buildings for the year was 2,606,104, an all-time record of attendance. This was an increase of 212,605 over the previous year’s attendance. April 1949 was the month of largest attendance with 371,871 visitors; August 1948, the second largest with 313,364. Records for the five buildings show the following number of visitors: Smithsonian, 494,880; Arts and In- dustries, 1,148,303; Natural History, 689,233; Aircraft, 198,648; Freer, 75,040. A summary of attendance records is given in table 1: TABLE 1.—Visttors to the Smithsonian buildings during the year ended June 80, 1949 Smith- Arts and Natural Adrorate Freer Year and month sonian Industries | History Buildin Gallery Total Building | Building | Building 8 of Art 1948 Duly a he oS ee eee 61, 529 128, 635 74, 243 24, 557 9, 510 298, 474 USUE ADC) a Say ee pe ee 65, 412 136, 704 75, 026 26, 672 9, 550 313, 364 September.a22 08-35 22 ee 45, 178 90, 321 61, 839 18, 460 7, 269 218, 067 Octobere-2)- 228s: fe Fast es 34, 460 66, 329 47, 962 13, 670 5, 460 167, 881 INovemberis= 2221 -iee ee 27, 380 50, 700 39, 829 11, 833 4,415 134, 157 December- 22s 222-22 22222-522 18, 242 42, 191 23, 419 8, 512 3, 153 95, 517 1949 JANUAIY See ee ee 26, 748 59, 837 37, 212 11, 085 4,124 139, 006 Mebruary 2 Seer aes See 22, 949 54, 470 35, 220 10, 842 4, 032 127, 513 Miarchste S220 eee ee 25, 650 66, 814 41, 452 12, 499 5, 092 151, 507 April see 32 ee eee 64, 804 177, 144 97,135 23, 532 9, 256 371, 871 VT yates oe Bate Mee non SOee ee 47, 718 142, 007 88, 029 19, 653 6, 172 303, 579 Jules. 5252222 s ses ean es 54, 810 133, 151 77, 867 17, 333 7, 007 290, 168 Total: === 442-5 3ee 494, 880 1, 148, 303 1689, 233 198, 648 75, 040 2, 606, 104 ' Not including 31,249 persons attending meetings after 4:30 p. m. SIXTEENTH JAMES ARTHUR ANNUAL LECTURE ON THE SUN In 1931 the Institution received a bequest from James Arthur, of New York, a part of the income from which was to be used for an annual lecture on some aspect of the study of the sun. SECRETARY’S REPORT 9 The sixteenth Arthur lecture was given in the auditorium of the National Museum on April 14, 1949, by Sir Harold Spencer Jones, Astronomer Royal of Great Britain, the arrangements being made through Dr. S. A. Mitchell, of the Leander McCormick Observ- atory, University of Virginia. The title of Sir Harold’s lecture was, “The Determination of Precise Time,” a subject on which he is a world authority. His lecture will be published in full in the Annual Report of the Board of Regents of the Smithsonian Institution for 1949. SUMMARY OF THE YEAR’S?ACTIVITIES OF THE BRANCHES OF THE INSTITUTION National Museum.—Approximately 446,000 specimens were added to the collections, for the most part as gifts or as transfers from Gov- ernment agencies, bringing the total number of catalog entries to 31,679,046. Outstanding accessions for the year included: In an- thropology, an important collection of 51 artifacts representing the work of American Indians, Eskimo of Alaska, and natives of Pacific islands, given by Georgetown University; 17 gold-embossed silver vessels given by the Government of Tibet to President Truman and in turn presented by him to the Smithsonian Institution; and valu- able skeletal remains recovered in northern Australia by Frank M. Setzler, a member of the Commonwealth of Australia-National Geo- graphic Society-Smithsonian Institution Expedition to Arnhem Land; in zoology, maramal specimens from many distant parts of the world including Northern Territory of Australia, Nepal, Malay Pen- insula, Korea, Okinawa, Philippine Islands, and New Guinea, 778 birds from Arnhem Land, Australia, and 1,164 from India and Nepal, 14,000 fishes from the Solomon Islands and the East Indies, and 5,000 from the Persian Gulf and the Red Sea; in botany, 2,382 plants of Fiji, 5,854 plants of Colombia, and 2,157 plants of China; in geology, 20 kinds of minerals hitherto unrepresented in the National collec- tions, a 42-carat brazilianite gemstone, the largest ever found in Brazil, an 8,750-gram stony meteorite that fell at Girgenti, Italy, and many thousands of fossil specimens collected by staff members in various parts of the United States; in engineering and industries, the original Wright Brothers’ aeroplane of 1903, a collection of elec- trical measuring instruments, early lamps, and electronic tubes, some of them made in the 1880’s, and an exhibit showing the development of electric hearing aids; in history, a group of relics bequeathed by Gen. John J. Pershing, including uniforms, flags, and medals, a note- worthy collection of European gold and silver coins from the four- teenth to the twentieth century presented by Paul A. Straub, of New York City, and a complete set of Allied military currency pre- sented by the Department of the Army. 10 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Field work was conducted in Arnhem Land in northern Australia, India and Nepal, the Persian Gulf and the Red Sea, New Zealand, the Canadian Arctic, nine different countries in South and Central America, and many parts of the United States. The Museum pub- lished its Annual Report, 3 Bulletins, 25 Proceedings papers, and 2 papers in the Contributions from the United States National Herbar- ium. The division of history was elevated to the status of a full department of the Museum, with five divisions—military history, naval history, civil history, numismatics, and philately. National Gallery of Art——During the year there were 1,529,568 visitors to the Gallery, an average daily attendance of 4,225. Acces- sions as gifts, loans, or deposits numbered 1,174, including 10 paint- ings and 50 prints and drawings from the estate of the late R. Horace Gallatin, and 891 prints and drawings from Lessing J. Rosenwald. Eleven special exhibitions were held at the Gallery, and two traveling exhibitions were circulated to art galleries, museums, and other organ- izations throughout the country. In response to inquiries received by the Gallery, nearly 1,000 research problems requiring reports were investigated, and advice was given regarding 233 works of art brought to the Gallery for opinion. Numerous books and articles on art sub- jects were published by staff members. New publications continued to be added to the literature available at the Gallery for purchase by the public. Some 15,000 persons attended the special tours of the Gallery, 20,000 the “Picture of the Week” talks, and 18,000 the lec- tures in the auditorium. The Gallery’s collections of art works has grown so fast that all available exhibition space was in use during the year. To provide for expansion, contracts have been let for the completion of 12 more galleries in unfinished areas of the Gallery building. ‘Some 50,000 persons attended the 46 Sunday evening concerts given in the Gallery’s East Garden Court. National Collection of Fine Arts.—At the annual meeting of the Smithsonian Art Commission of December 7, 1948, a number of paintings were accepted for the National Collection. The Commis- sion passed a resolution calling attention to the inadequacy of the present art exhibition facilities in the National Museum and recom- mending that the Secretary of the Smithsonian Institution take action to provide proper space for the preservation and exhibition to the public of the National Collection of Fine Arts. Two miniatures were acquired through the Catherine Walden Myer fund. Under the provisions of the Ranger bequest, seven paintings temporarily as- signed to various art institutions were recalled for final consideration by the Smithsonian Art Commission. Two of these paintings were accepted for the National Collection, and the others were returned to the institutions to which they were originally assigned. A large SECRETARY'S REPORT 11 amount of information on art subjects was furnished to visitors in person, as well as by mail and phone. Members of the staff lectured on art topics to several organizations, and six special art exhibitions were held during the year, for most of which catalogs were furnished by the organizations sponsoring the exhibitions. Freer Gallery of Art.—Additions to the collections included Chinese bronze, jade, lacquer, marble, and painting; Syrian glass; Syrian or Egyptian gold; Arabic manuscript; Persian manuscript, painting, and stone sculpture; Indian painting; and Turkish painting. The work of the professional staff was devoted to the study of new accessions and to research within the collection of Chinese, Japanese, Iranian, Arabic, and Indian materials. Reports were made upon 2,563 objects and 372 photographs of objects submitted to the Gallery for examina- tion, and 369 Oriental language inscriptions were translated. The repair and restoration of the walls of Whistler’s Peacock Room were completed early in the year, and work was begun on the ceiling. Visitors to the Gallery numbered 74,846 for the year, and 1,724 came to the Gallery offices for special purposes. Sixteen groups were given instruction in the exhibition galleries by staff members, and 13 lec- tures were given in art galleries and museums, before clubs, and to various associations. Bureau of American Ethnology.—Dr. M. W. Stirling, Director of the Bureau, devoted 4 months to a continuation of his archeological work in Panamé in cooperation with the National Geographic Society. Heretofore undescribed ceramic cultures were found at Utivé and Barriles, and much new information was obtained on the classic Chiriqui and Veraguas cultures. Dr. Frank H. H. Roberts, Jr., contmued to direct from Washington the very extensive operations of the River Basin Surveys, a unit of the Bureau created to rescue important archeological sites threatened by the construction of dams and the creation of river basin reservoirs. ‘The work was done in cooperation with the National Park Service, the Bureau of Reclama- tion, the Army Corps of Engineers, and local organizations. Surveys of threatened sites covered 69 reservoir areas in 21 States. Since the program started, 2,107 archeological sites have been located and recorded, and of these, 456 have been recommended for excavation or testing before they are destroyed by construction work. Dr. John P. Harrington continued his revision of the Maya grammar. Toward the end of the year he went to Old Town, Maine, to pursue ethnological and linguistic studies on the Abnaki Indians. Dr. Henry B. Collins, Jr., conducted archeological excavations at Frobisher Bay on Baffin Island in the Canadian Arctic. Ruins were found of old Eskimo semisubterranean houses made of stones, whale bones, and turf, the evidence showing that the site has been occupied succes- 12 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 sively by Eskimos of both the prehistoric Dorset and Thule cultures. Dr. William N. Fenton continued his field work and library research on the Iroquois Jndians, obtaining the life history of an aged Seneca and recording Seneca rituals, prayers, and legends. Dr. Gordon R. Willey devoted the year to studying and writing up the results of previous field work. His monographic work, “Archeology of the Florida Gulf Coast,”’ was completed and at the close of the year was in process of being published by the Smithsonian Institution. The Institute of Social Anthropology, an autonomous unit of the Bureau, is financed by State Department funds to carry out coopera- tive training in anthropological teaching and research with the other American republics. Institute staff members, under the directorship of Dr. George M. Foster, Jr., continued to give courses in anthropology and to conduct cooperative research and field work in Brazil, Colom- bia, México, and Pert. The Bureau published its Annual Report and two Publications of the Institute of Social Anthropology. The last two volumes of the Handbook of South American Indians, volumes 5 and 6, were in press at the close of the year. International Hachanges.—The Smithsonian International Exchange Service is the official United States agency for the interchange of gov- ernmental and scientific publications between this country and the other nations of the earth. ‘The Exchange Service handled during the year a total of 840,125 packages of publications, weighing 796,700 pounds. These figures represent an increase over the previous year of 80,006 packages, but a decrease of 15,489 pounds in weight, indicat- ing by the lighter weight per package that most institut'ons have about completed shipment of material held up during the war. Shipments are now made to all countries except Rumania, and efforts to resume exchanges with that country are being continued. The number of sets of United States official publications sent abroad in exchange for similar publications of other countries is now 96—58 full and 38 partial sets. There are also sent abroad through the Exchange Service 81 copies of the Federal Register and 75 copies of the Con- gressional Record. National Zoological Park.—The collection was improved during the year by the addition of a number of rare animals. At the close of the fiscal year there were 3,724 specimens in the collection, an increase of 927 over the previous year. These represented 755 different species, an increase of 65. Among the rare or unusual animals received by gift, exchange, or purchase were the rare Meller’s chameleon, a spectacled bear, a pair of pigmy marmosets—smallest of all monkeys, an African two-horned rhinoceros, a pair of wombats, a pigmy ant- eater, orang-utans, and chimpanzees. The total number of creatures SECRETARY'S REPORT 1433 born or hatched at the Zoo was 157—56 mammals, 62 birds, and 39 reptiles. Personne! recruitment and training for the organization progressed satisfactorily, and the most needed repairs and minor im- provements to buildings and grounds were carried out. The year’s total of visitors to the Zoo was the largest ever recorded—3,346,050, an increase of more than 300,000 over the previous year. Groups from schools, some as far away as Maine, Florida, Texas, and Cali- fornia, numbered 1,844, aggregating 93,632 individuals. Astrophysical Observatory —Year-long tests at the three most promising sites for a new high-altitude solar observing station indicate that the best skies prevail at the Clark Mountain, Calif., site, the second-best site being Pohakuola, Hawaii. Estimates of the cost of establishing a field station on Clark Mountain, however, proved to be in excess of available funds, forcing postponement of building opera- tions. Data and tables were prepared which simplify computations at the field observing stations by eliminating the tedious curve-plotting process heretofore used in obtaining the air mass. Daily observations of the solar constant of radiation were continued at the Montezuma, Chile, and Table Mountain, Calif., stations. Intercomparisons between the substandard silver-disk pyrheliometer S. I. No. 5 and the instruments in use at Miami, Montezuma, and Table Mountain show no material changes in constants, confirming the adopted scale of pyrheliometry. Special radiation measurements started in 1945 at Camp Lee, Va., under contract with the Office of the Quartermaster General, were continued there, half of the year by the Observatory and half by the Quartermaster Board; similar measurements were also made at Miami, Fla., and at Montezuma, Chile. The work of the Division of Radiation and Organisms has been concerned chiefly with reorganizing and reequipping the laboratories. Besides new office space which has been established, five rooms are being converted into constant-condition rooms for biological experimentation, and four chemistry laboratories will be available. In addition, a photographic laboratory, an X-ray room, a cytology laboratory, an electronics laboratory, and two general laboratories are being arranged. National Air Museum.—The Air Museum was given the responsi- bility of receiving, bringing to Washington, and preparing for exhibi- tion the original Wright Brothers aeroplane of 1903, presented to the National Museum in December 1948. A storage depot to be used by the Air Museum until it has a building of its own was acquired in November 1948 in the former Douglas Aircraft plant at Park Ridge, Ill. There a field organization was installed, and the Air Museum assumed custody of the storage facility itself and the large collection of planes and other aeronautical material stored there by the U. S. Air Force for the Museum. The Advisory Board held three meetings 14 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 during the year which were devoted mainly to advancing the acquisi- tion of a building site and a suitable museum building in the Wash- ington area. The Museum expects during the coming year to sub- mit to Congress a report regarding sites and a building, the prelim- inary study of which has been prepared in cooperation with the Public Buildings Administration. Among outstanding accessions of the year were the Swoose, the historic B~17—D bomber that served through- out World War II from Bataan to the defeat of Japan, presented by the city of Los Angeles; Maj. Alford Williams’ renowned Gulfhawk-2 presented by the Gulf Oil Co.; a Japanese Baka Bomb, or “suicide plane,” transferred by the Department of the Navy; and 10 scale models of recent types of Naval aircraft received from the manufac- turers who produced the original planes. New accessions totaled 122 objects from 40 different sources. Canal Zone Biological Area.—A new building for woodworking and carpenter shops and for living quarters for the warden-caretaker was completed during the year, the old quarters being converted into a two-room laboratory unit. Work on the new 14,000-gallon water tank was halted by heavy rains but can be completed with 2 or 3 weeks of dry weather. The most urgent needs are the fireproofing of existing buildings and the construction of a new six-room laboratory and storage building. Twenty-nine scientists representing many different organizations worked at the laboratory during the year, and their contributions have added materially to our knowledge of tropical life. Among the interesting researches were the work of Drs. Scho- lander and Walters of the Arctic Research Laboratory at Point Barrow, Alaska, on the metabolic reactions to temperature in various animals and plants in order to obtain a tropical counterpart for similar work on Arctic forms in Alaska; the studies of Drs. Clark and Soper of the Research Laboratory of Eastman Kodak on the effects of tropical conditions on photographic equipment and materials, including color film; and the Resident Manager’s own special studies, particularly the long-term termite-resistance tests. PUBLICATIONS In carrying out the diffusion of knowledge, the Institution issues eight regular series of publications and six others that appear less fre- quently. All these series, embodying the results of Smithsonian researches, are distributed free to more than a thousand libraries, both here and abroad, as well as to a large list of educational and scientific organizations. The findings of Smithsonian scientists, chiefly in the fields of anthropology, biology, geology, and astrophysics, are therefore made readily available to all through this wide free distribution. SECRETARY'S REPORT 15 A total of 71 separate volumes and pamphlets were issued during the past year. Among the outstanding publications to appear were Dr. Henry Field’s compilation in the Smithsonian Miscellaneous Collec- tions entitled ‘Contributions to the Anthropology of the Soviet Union,”’ which presents, for the first time in English, accounts of recent findings in this little-known area; a revised edition of the popu- lar handbook of the National Aircraft Collection, which is in effect a brief history of aeronautics from the mythical flying horses of antiquity to the supersonic jet planes of today; two more volumes in the famous series of Life Histories of North American Birds, prepared by A. C. Bent, bringing to 17 the number so far issued in the series; and a paper by Jason R. Swallen on new grasses from several countries of South and Central America, in the Contributions from the United States National Herbarium. The total number of copies of publications in all series distributed during the year was 267,491. A complete list of the year’s publica- tions will be found in the report of the Chief of the Editorial Division, Appendix 12. LIBRARY Of the 57,671 publications received by the Smithsonian library during the year, 7,287 came as gifts from many different donors. Another 17,713 were periodicals mostly received in exchange for Smithsonian publications from research institutions and other scien- tific and educational organizations throughout the world. Containing the record of progress in science and technology, these periodicals are indispensable in the prosecution of the Institution’s own work. Increasingly heavy demands upon reading and reference services of the library were noted during the year, the interlibrary loans total- ing 2,619 publications to 89 different libraries. The new position of assistant librarian in charge of the Astrophysical Observatory library was filled by the promotion of an acquisitions assistant. New exchanges arranged during the year numbered 338; 6,884 volumes and pamphlets were cataloged, and 31,184 cards were added to catalogs and shelflists; 1,060 volumes were sent to the bindery, and 1,026 were repaired in the Museum. At the close of the year, the library’s holdings totaled 921,206 volumes, more than half of which are housed in the Library of Congress as the Smithsonian Deposit. Respectfully submitted. ALEXANDER WETMORE, Secretary. APPENDIX 1 REPORT ON THE UNITED STATES NATIONAL MUSEUM Srr: I have the honor to submit the following report on the condi- tion and operations of the United States National Museum for the fiscal year ended June 30, 1949. COLLECTIONS Approximately 446,000 specimens (88,000 less than last year) were incorporated into the National collections during the year and were distributed among the six departments as follows: Anthropology, 4,099; zoology, 279,621; botany, 38,708; geology, 109,499; engineering and industries, 2,610; and history, 11,104. The decrease in the number of specimens accepted for the Museum’s collections may be attributed in part to the inadequacy of available storage facilities for the preser- vation of such materials; consequently, a finer screening of collections from prospective donors is now mandatory. Most of the accessions were acquired as gifts from individuals or as transfers from Govern- ment departments and agencies. The complete report on the Museum, published as a separate document, includes a detailed list of the year’s acquisitions, of which the more important are summarized below. Catalog entries in all departments now total 31,679,046.! Anthropology.—The most noteworthy additions to the archeological collections were as follows: A black-figured Attic lecythus of the fifth century, B. C., presented to President Harry S. Truman as a token of gratitude from the people of Greece and lent by the President; 11 gold-plated ornaments from Veraguas, Panam4, and 2 gold fishhooks from Colombia, a gift from Karl P. Curtis; and 47 prehistoric earthen- ware vessels from the Valley of Nasca, Peri, presented to the late Gen. John J. Pershing by former Peruvian President Augusto B. Leguia and donated by General John J. Pershing. Handicrafts and material culture of many of the world’s peoples were represented in the additions to the ethnological collections. An unusually important collection of 51 specimens representing the work of American tribes of the Great Plains and the Great Lakes, of Arizona and New Mexico, as well as of the Eskimo of Alaska, of the Igorot of the Philippine Islands, and of the Marquesans and Maori of the 1 The revised tabulation of the National collection of insects during the past year, in addition to the normal increment, has increased last year’s total by more than 4,400,000 specimens. 16 SECRETARY'S REPORT 17 Southeast Pacific, assembled over a period of more than a century, was presented by Georgetown University. President Harry 8S. Tru- man presented to the Smithsonian Institution 17 gold-embossed silver vessels received at the White House as a gift from the Government of Tibet in appreciation of an American gift of wireless receiving and transmitting sets made during World War II. Included are two butter lamps and stands, four teacup stands and covers, two bowls for grain offerings, one teapot, and two beer mugs, all decorated in gold-embossed designs derived from Chinese-Tibetan folklore and Buddhist religious art. A collection of 287 folk, costume, and his- torical portrait dolls, representing the native dress of peoples of many lands, was received as a bequest from the late Mrs. Frank Brett Noyes. The Don Diego Columbus mahogany table, traditionally known as the writing desk of Diego Columbus, was conditionally bequeathed by Mrs. Edith Keyes Benton. This table had been preserved for centuries in the cathedral of Santo Domingo City and was presented by Archbishop Nouel to Commander Frederick L. Benton, U.S.N., in recognition of his work in Santo Domingo during the influenza epidemic of 1918. One of the rarest of musical instruments, a musical gong, kyung, carved from white marble, was presented by Ju Whan Lee, director of the Korean Court Music Conservatory at Seoul, Korea. The largest accession received by the division of physical anthro- pology consisted of the skeletal remains recovered in northern Australia by Frank M. Setzler, a member of the Commonwealth of Australia- National Geographic Society-Smithsonian Institution Expedition to Arnhem Land. Australian skeletal material available for study in the United States is rather limited. Four casts of African fossil primates, which illustrate certain characteristics of antecedent special- ization, were also acquired during the year. Zoology.—The collections made by the Museum staff detailed to the Arnhem Land field expedition, under the joint sponsorship of the Commonwealth of Australia, National Geographic Society, and the Smithsonian Institution, have added many previously unrepresented forms of animal life to the National collections. These collections included not only vertebrates but invertebrates as well. Accessions that enhanced the usefulness of the mammalian collec- tion came from the Northern Territory of Australia, Nepal, Malay Peninsula, Korea, Okinawa, Philippine Islands, New Guinea, and New Hampshire. Field work financed wholly or in part by the W. L. Abbott fund resulted in the addition of birds not hitherto represented in the National collection. Included among these accessions were 2,815 skins and 388 eggs of Colombian birds; 900 skins, 24 skeletons, and 2 sets of eggs of Panamanian birds; 778 bird skins, many of which were not represented in the collection, as well as 51 skeletons and 2 eggs 18 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 from Arnhem Land, Australia; and 1,164 bird skins procured by the joint National Geographic Society-Yale University-Smithsonian Institution Expedition to India and Nepal. Other accessions com- prised 611 bird skins from Nyasaland: 177 birds and 1 egg from northeastern Venezuela: 171 bird skins from Pacific War areas; and 125 bird skins from Korea. Snakes, lizards, and frogs from Arnhem Land, amphibians from Perit, reptiles and amphibians from Honduras, and a general collection from Virginia and North Carolina constituted the most important additions to the herpetological collection. The most noteworthy accessions received by the division of fishes were nearly 14,000 specimens from the Solomon Islands and the East Indies, which were presented by Dr. Wilbert M. Chapman; 14,300 from Arnhem Land; and approximately 5,000 from the Persian Gulf and the Red Sea, resulting from a survey sponsored by the Arabian- American Oil Co. Other important collections of fishes came from Puerto Rico, Panam4, British Columbia, and Florida. Approximately 25,000 miscellaneous insects from South Pacific Islands came to the Museum by transfer from the U. S. Commercial Co. Among other large lots were approximately 12,000 flies; 3,500 chalcidoid wasps; 500 beetles; and some 53,000 insects transferred from the United States Bureau of Entomology and Plant Quarantine. During the year considerable significant material was added to the marine invertebrate collection, of which the most important accessions were 11,765 miscellaneous invertebrates from the Department of Zoology, University of California; 70 lots of paratypes, hypotypes, and topotypes of hydroids from the Allan Hancock Foundation, University of Southern California; 760 marine invertebrates from California and Mexice; 709 specimens from Bahama Islands; 1,781 from Pacific Islands and California; 452 from the Persian Gulf and the Red Sea; and 859 from Arnhem Land. By transfer from the Office of Naval Research, the Museum acquired 3,668 invertebrates from Point Barrow, Alaska. The United States Geological Survey transferred 568 specimens from the Marianas Islands. A rare deep-water Pleurotomaria, dredged at a depth of 160 fathoms off Natal, South Africa, and presented by Dr. Cecil von Bonde, con- stituted the most notable accession received by the division of mollusks. From other sources the division received 250 Peruvian terrestrial and fresh-water mollusks and 540 marine mollusks from Canton Island, and 150 Japanese land mollusks. Exchanges brought to the Museum approximately 1,080 shells from Spain and lesser numbers from South Africa, Italy, and Cuba. By transfer the Museum received about 1,200 mollusks obtained in the Caroline Islands from the United States Geological Survey; approximately 30,600 specimens SECRETARY’S REPORT 19 from the Naval Medical Research Institute; and 600 marine and land shells of the Solomon Islands from the Naval Medical School. Mem- bers of the staff obtained about 1,200 mollusks in Arnhem Land and some 1,500 in the region of the Persian Gulf and the Red Sea. Botany.—As exchanges, the National Herbarium received 2,382 plants, comprising a collection made in Fiji by Dr. A. C. Smith, from the Arnold Arboretum of Harvard University; 5,854 plants of Colom- bia from the Facultad de Agronomfa, Universidad Nacional, Medellin; and 2,157 Chinese plants from the National Szechwan University. The Division of Rubber Plant Investigations, United States Depart- ment of Agriculture, transferred 865 plants from eastern Colombia. The Oficina de Estudios Especiales, Mexico City, presented 394 Mexican grasses. = nc ne ae ne Landscape with Boatman. A painting by Murillo, “The Return of the Prodigal Son,” given by the Avalon Foundation, was accepted by the Board of Trustees on December 10, 1948. At the same time the Board accepted the portrait of Daniel Boardman, by Ralph Earl, from Mrs. W. Murray Crane; “Interior of a Church,” by Pieter Neeffs, from Senator Theo- dore Francis Green of Rhode Island; and two paintings, ‘‘Repose,”’ by John Singer Sargent, and “Head of a Girl,” by James Abbott McNeill Whistler, from Curt H. Reisinger. On December 22, 1948, the Board of Trustees accepted from Dr. G. H. A. Clowes a painting, “Allegory,” Venetian School about 1500, and from Vladimir Horo- witz a painting, ““Head of a Young Girl,” by Renoir. The Board of Trustees accepted from Miss Georgia O’Keeffe on March 8, 1949, a gift of the following three paintings: Artist Title Marsden Hartley...) =) 322 al 2 es Se Landscape No. 5. Ar Chun GAM ONG 55-285 ee Sane Seeue ess Moth Dance. Georgia. Osieetiens. rere oe Ae ee To be selected later. 28 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 During the fiscal year, the portrait of Captain Patrick Miller by Raeburn, previously on loan, was given to the Gallery by Mrs. Dwight Davis. SCULPTURE On December 10, 1948, the Board of Trustees accepted from Stanley Mortimer, Jr., a ‘“‘Portrait Bust of a Member of the Order of San Iago” attributed to Leone Leoni, which had previously been on loan to the Gallery. At the same time the Board accepted from Miss Mildred Howells a portrait medallion of her father, William Dean Howells, and herself, by Augustus Saint-Gaudens, to be held for a National Portrait Gallery. PRINTS AND DRAWINGS A gift from Lessing J. Rosenwald of 309 additional prints and drawings was accepted on December 10, 1948, to be added to the Lessing J. Rosenwald Collection. At the same time, two volumes of “The Georgics’”’ of Virgil with 119 illustrations by Andre Dunoyer de Segonzac were accepted as a gift from the artist. This gift was inspired by an earlier gift to the Gallery of a collection of Segonzac’s prints and drawings made in memory of the late Frank Crowninshield. The Board of Trustees, during the fiscal year, received 50 prints and drawings from the collection of the late R. Horace Gallatin. On March 8, 1949, the Board accepted from Miss Georgia O’ Keeffe three water colors by John Marin entitled “Movement, Boat and Sea, Deer Isle, Maine,’ “White Mountain Country, Summer,” and “Storm over Taos, New Mexico.”’ The Board of Trustees accepted from Mr. Rosenwald on May 3, 1949, 582 additional prints and draw- ings. Received during the fiscal year from George Matthew Adams were 20 etchings by Alphonse Legros. PHOTOGRAPHS The Board of Trustees on March 8, 1949, accepted from Miss Georgia O’Keeffe a key set of photographs, consisting of about 1,500 prints, by Alfred Stieglitz. EXCHANGE OF WORKS OF ART During the fiscal year 1949 the Board accepted the offer of Chester Dale to exchange the portrait of Ralph Waldo Emerson by Sully, which was being held for the National Portrait Gallery, for the por- trait of the Sicard David Children by Sully, which was then on loan to the Gallery. The Board also accepted the offer of Lessing J. Rosenwald to exchange the prints ‘Sacrifice to Priapus,”’ by Jacopo de Barbari, ‘Conversion of St. Paul,’’ by Lucas van Leyden, and SECRETARY’S REPORT 29 “Solomon Worshipping Idols,’ by the Master M. Z., for superior impressions of like prints now included in the Rosenwald Collection at the National Gallery of Art. WORKS OF ART ON LOAN During the fiscal year 1949 the following works of art were received on loan by the National Gallery of Art: From Artist Chester Dale, New York, N. Y.: 1 BCR (0) oes ae de Ag ee ire oN ee Pee Domergue. Mrs bhilipyliydics 8-2 ss 2 seo seee eee S Zuloaga. SanvSepulvedaase= = sss te ee eee Zuloaga. iarRubiaydelAbanicoss222-- 2-2 sse— ee osa2 Zuloaga. Mrs. Brooks Goddard, Paris, France (via the National Collection of Fine Arts): Mrursical eins pina tio epee eae rere Romaine Brooks Goddard. PEHewBaleory 55 22 5 UM A sere es Se eg Romaine Brooks Goddard. Sketcne fete ee ssf oe ee ey, Bele eee Romaine Brooks Goddard. SelfsPortral tees eee ee eee ee eee Romaine Brooks Goddard. Alfred Stieglitz Collection: (Miss Georgia O’ Keeffe, New York, N. Y.) Chimneys and Water Tower-------------- Demuth. AGCowssiskulliwithpRediers= eee O’ Keeffe. Ihinerand=Curviess=s28s8s cose ee eee eee O’ Keeffe. Chauncey Stillman, New York, N. Y.: AVHalberdier 24 tee ee ss ee ae asl Pontormo. George Matthew Adams, New York, N. Y.: Sretchings an 2 ole as sue ee ee awe Alphonse Legros. C. S. Gulbenkian, Lisbon, Portugal: 28 pieces of Egyptian sculpture. 3 pieces of eighteenth-century French fur- niture. 1 fourteenth-century Arabian bottle. 1 sixteenth-century Persian rug. 7 eighteenth-century French books. The Italian Government: AVmarblerscatueOi Davide sane sees ae ee Michelangelo. Robert Woods Bliss, Washington, D. C.: 32 objects of Pre-Columbian art. LOANED WORKS OF ART RETURNED The following works of art on loan were returned during the fiscal year 1949: To Artist Chester Dale, New York, N. Y.: OnitherBeachee a ane ee eee eae ees Winslow Homer. Mme. Charlotte Fuerstenberg, at New York, INceyY -: Sea abr Pstaquers. sess o2— oneness Cezanne. 30 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Anonymous loan: Paradise, Valleyi43t 2-442 42-63-20 ee en John La Farge. Robert Woods Bliss, Washington, D. C.: 16 objects of Pre-Columbian art. Peabody Museum, Harvard University, Cam- bridge, Mass.: 70 objects of Pre-Columbian art. WORKS OF ART LOANED During the fiscal year 1949, the Gallery loaned the following works of art for exhibition purposes: To Artist Albright Art Gallery, Buffalo, N. Y.: Joseph; Widener. -.-.---2 See +s. sseesS Augustus John. Art Institute of Chicago, Chicago, IL: Alexander: Hannltonassse oo eo Trumbull. William) Dhormtone -enAs) 22-2 ee ee Stuart. Columbus Gallery of Fine Arts, Columbus, Ohio: iAbrahsimeltineoinivres 62 2S. sano ee Healy. Corcoran Gallery of Art, Washington, D. C.: ‘The White Girl. 2-2 sa.=0* = ess eee Whistler. Dallas Museum of Fine Arts, Dallas, Tex.: George Washington (Vaughan-Sinclair) ----- Stuart. Dayton Art Institute, Dayton, Ohio: ackawanna Valleys sof ses- 22. 2 eee Inness. Fort Worth Art Association, Forth Worth, Tex.: Breezing Wp. eee aA a ee Winslow Homer. Metropolitan Museum of Art, New York, N. Y.: Captain Charles Stewart. -.--.-.--------- Sully. Pack Memorial Library, Asheville, N. C.: Tomas *Vawsone 26 sa 2se ese = = ee Mather Brown. Henry, Laurense... 2 = 23st atest est Copley. Andre wa acCkSON= =a = esac arene eee eee Earl. WiilltaminatVloores ae ene ee = = ae Feke. General) William: Moultries === 22-- — = === Charles Willson Peale. JohnmiC: Calhoune ses. 226-2. ae Rembrandt Peale. JOUnNeBaptista Ashe nas sas ee ee eee Stuart. Matilda Caroline; @ruger. = .=..-----=-—-=- Stuart. Hrancis, Hopkinson... 25 S22_2- = 22 ee Sully. Annebiddle Hopkinson] —5-—. 222-4... 2 355-2 Sully. JosiasrAllston ease oe ae eee eee Theus. Walliam Rogerst=.4 ies ease es ee eee Trumbull. Portraits, Inc., New York, N. Y.: Mrs#Chester Dale: si) 210 0 sees fat or Bellows. Mr Chester Dale. 25. == 252-2. GASES Bellows. Scott and Fowles, New York, N. Y.: Joseph Widener... 2.522.225. 5.css5558 Augustus John. SECRETARY'S REPORT oil EXHIBITIONS During the fiscal year 1949 the following exhibitions were held at the National Gallery of Art: American Paintings from the Collection of the National Gallery of Art. Exhi- bition of American paintings, featuring a group of portraits from Pocahontas to General Eisenhower. Continued from previous fiscal year, through July 11, 1948. American Folk Art. Exhibition consisting of 104 water-color renderings from the Index of American Design. July 18 to September 7, 1948. American Graphic Art from the Eighteenth Century to the Present Day. Selection from the collections of the Library of Congress, the Smithsonian Institu- tion, and the National Gallery of Art. September 19 to November 14, 1948. Paris the Favorable Climate. Exhibition of prints and drawings by Bonnard, Vuillard, Maurice Denis, Andre Dunoyer de Segonzac, and Matisse, arranged in memory of Frank Crowninshield. November 21, 1948, to January 11, 1949. Michelangelo’s ‘‘David.’’ Lent to the National Gallery of Art by the Italian Government. January 24 to June 28, 1949. Gulbenkian Collection of Egyptian Sculpture. Lent for an indefinite period to the National Gallery of Art by C. 8. Gulbenkian. Opened January 30, 1949. Studies of Medieval Cathedrals. Exhibition of photographic studies lent to the National Gallery of Art by Clarence Ward, head of the Department of Fine Arts, Oberlin College. January 30 to February 13, 1949. Gulbenkian Collection of Eighteenth Century French Objects. Additions to earlier loan by C. S. Gulbenkian, on exhibition at the National Gallery of Art for an indefinite period. Opened February 20, 1949. American Paintings from the Collection of the National Gallery of Art. Feb- ruary 20 to April 10, 1949. Early Italian Engraving. Exhibition of early Halian engravings, lent to the National Gallery of Art by various museums and anonymous lenders. April 17 to June 19, 1949. R. Horace Gallatin Collection. Exhibition of prints bequeathed to the National Gallery of Art by Mr. Gallatin. Opened June 26, 1949. The following exhibitions were displayed in the cafeteria corridor of the National Gallery of Art during the fiscal year 1949: Whistler Prints. Rosenwald Collection; one gift of Myron A. Hofer. Con- tinued from previous fiscal year through July 18, 1948. Audubon Prints. Mrs. Walter B. James Collection. July 20 to December 12, 1948. Index of American Design. Water-color renderings of early American toys. December 13, 1948, to February 15, 1949. Index of American Design. Water-color renderings of early American furniture and textiles. February 16 to March 28, 1949. Legros Prints. George Matthew Adams Collection. March 29 to May 15, 1949. Seymour Haden Prints. Rosenwald Collection and gift of Miss Elisabeth Achelis. May 16 to June 12, 1949. Ostade Prints. Rosenwald Collection and gift of Mrs. Addie Burr Clark. Opened June 13, 1949. a2 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 TRAVELING EXHIBITIONS Rosenwald Collection.—Special exhibitions of prints from the Rosen- wald Collection were circulated to the following places during the fiscal year 1949: Kenneth Taylor Galleries, Nantucket, Mass.: 26 French prints. July 26 to August 23, 1948. Watkins Gallery, American University, Washington, D. C.: 26 French prints. October 13 to 30, 1948. Los Angeles County Museum, Los Angeles, Calif.: 20 Blake prints. October 1948. Wyncote Woman’s Club, Wyncote, Pa.: 11 prints. October 17 to 23, 1948. Rutgers College, New Brunswick, N. J.: 9 Italian prints. October 1948. Fogg Museum of Art, Harvard University, Cambridge, Mass.: 1 Rembrandt drawing. November 1948. Museum of Modern Art, New York, N. Y.: 1 Munch print. November 1948 to January 1949. Walters Art Gallery, Baltimore, Md.: 6 Gavarni drawings. January 22 to March 6, 1949. Walters Art Gallery, Baltimore, Md.: 5 miniatures. January 27 to March 138, 1949. City Art Museum, St. Louis, Mo.: 17 prints. March 1949. Institute of Contemporary Arts, Washington, D. C.: 11 Klee prints. March 21 to April 22, 1949. Philadelphia Museum of Art, Philadelphia, Pa.: 3 Lehmbruck prints. May 1949. Art Gallery of Toronto, Toronto, Canada: 67 prints. May 1949. SECRETARY'S REPORT 33 Index of American Design.—During the fiscal year 1949 exhibitions from this collection were shown at the following places: Library of Congress, Washington, D. C. Western Reserve Historical Society, Cleveland, Ohio. Shaker Village Work Camp, Pittsfield, Mass. New York State Historical Association, Cooperstown, N. Y. Damariscotta Information Bureau, Damariscotta, Maine. University of ‘Tennessee, Tenn. Wustum Museum of Fine Arts, Racine, Wis. City Art Museum, St. Louis, Mo. William Rockhill Nelson Gallery, Kan- sas City, Mo. Munson- Williams-Proctor Institute, Utica, N. Y. Toledo Museum of Art, Toledo, Ohio. Mint Museum, Charlotte, N. C. Museum of Fine Arts, Montgomery, Ala. Knoxville, Schenectady Museum, Schenectady, Na ¥2 University of Oklahoma, Norman, Okla. University of Michigan, Ann Arbor, Mich. North Carolina College, Durham, N. C. Art Institute, Zanesville, Ohio. Atlanta University, Atlanta, Ga. Currier Gallery of Art, Manchester, N. H. Stephens College, Columbia, Mo. Brown University, Providence, R. I. Fort Valley State College, Fort Valley, Ga. Washington College, Chestertown, Md. Everhart Museum, Scranton, Pa. Art Gallery, Grand Rapids, Mich. Florida Agricultural and Mechanical College, Tallahassee, Fla. Farnsworth Museum, Rockland, Maine. Tuskegee Institute, Tuskegee, Ala. Young Playways, Inc., Washington, Dr CE: Smith College, Northampton, Mass. Prairie View University, Prairie View, Tex. University of North Dakota, Grand Forks, N. Dak. American University, Washington, D. C. Rockford Art Association, Rockford, II. Sweet Briar College, Sweet Briar, Va. Arkansas Agricultural, Mechanical and Normal College, Pine Bluff, Ark. Alfred University, Alfred, N. Y. Fisk University, Nashville, Tenn. St. Paul Public Library, St. Paul, Minn. Spelman College, Atlanta, Ga. Arnot Art Gallery, Elmira, N. Y. Kenneth Taylor Galleries, Nantucket, Mass. CURATORIAL ACTIVITIES The Curatorial Department accessioned 1,118 new gifts to the Gallery during the fiscal year. Advice was given in the case of 233 works of art brought to the Gallery for opinion, and 58 visits were made by members of the staff in connection with proffered works of art. Almost 1,000 research problems requiring reports were investigated in response to inquiries received by the Gallery. During the year, 16 individual lectures were given by members of the curatorial staff, both at the Gallery and elsewhere. In addition Miss Elizabeth Mongan gave a seminar at Alverthorpe, Jenkintown, Pa., for Swarth- 34 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 more College honor students; Charles Seymour, Jr., gave a course at Johns Hopkins University on Renaissance Art; and Charles M. Richards gave a survey course on art history under the auspices of the Department of Agriculture. Miss Mongan also made the arrangements for Arthur M. Hind’s American lecture tour, in connec- tion with the publication of Part II of his ‘‘Harly Italian Engraving,” under Gallery auspices. Mr. Seymour served on three and Miss Mongan on two art juries. Special installations were prepared for: the Michelangelo “David” lent to the National Gallery of Art through the courtesy of the Italian Government; 28 pieces of Egyptian sculpture lent to the Gallery by C. S. Gulbenkian placed on exhibition in January 1949; and eighteenth-century furniture and books also lent by Mr. Gulben- kian. The cataloging and filing of photographs in the George Martin Richter Archive continued to make progress, with the gradual enlargement of the collection. Further activities of the department are indicated under the heading of “‘ Publications.” RESTORATION AND REPAIR OF WORKS OF ART Necessary restoration and repair of works of art in the Gallery’s collections were made by Stephen S. Pichetto, Consultant Restorer to the Gallery, until his death in January 1949. No successor to Mr. Pichetto has as yet been appointed, but necessary minor repairs on the works of art have been continued under the care of Mr. Pichetto’s residual staff. All work was completed in the Restorer’s studio in the Gallery, with the exception of the restoration of two paintings, work on which is being completed in the New York studio of S. S. Pichetto, Inc. PUBLICATIONS During the year Mr. Cairns published two books, ‘‘The Limits of Art,’’? Pantheon Books, Inc., and ‘‘Legal Philosophy from Plato to Hegel,” Johns Hopkins Press. He also edited a volume entitled “Lectures in Criticism,’”’ Pantheon Books, Inc., and contributed an introduction to “Epicurus, My Master,’’ by Max Radin, University of North Carolina Press. He also contributed articles and reviews to the Columbia Law Review, Human Events, Saturday Review of Literature, New York Herald Tribune, Baltimore Evening Sun, Law and Contemporary Problems, The Scientific Monthly, and to the volume El Actual Pensamiento Juridico de los Estados Unidos, Buenos Aires. A series of 12 articles on masterpieces in the Gallery, prefaced by one entitled ‘‘New Friends for Old Masters,” is being published by John Walker in the Ladies Home Journal. An article by Mr. SECRETARY’S REPORT 35 Walker, “The Art of Duplicating Great Art,” appeared in Vogue on August 15, 1948, and another, ‘‘American Masters in the National Gallery,” in the National Geographic Magazine in September 1948. Mr. Walker also contributed two book reviews, the first reviewing Bernard Berenson’s ‘‘ Aesthetics and History in the Visual Arts” to the October 1948 Gazette des Beaux-Arts, and the second, entitled ‘The Philosophy of a Connoisseur,’ a review of Mr. Berenson’s ‘Sketch for a Self-Portrait,’ to the New York Times for April 24, 1949. Charles Seymour, Jr., published two articles, ““Note on the Relationship between an Illustration by Travies de Villers and Daumier’s ‘Le Fardeau’,” in the Journal of the Walters Gallery for 1948, and in the Summer Bulletin of the Columbus Gallery of Fine Arts the text of the address given by him for the inauguration of a group of sculpture by Georg Ehrlich in the Columbus Gallery of Fine Arts. Printing of ‘‘Masterpieces of Sculpture from the National Gallery of Art,’ a volume prepared by Mr. Seymour, was begun during the summer of 1949. Mrs. Fern R. Shapley has written two book reviews, a review of Bernard Berenson’s ‘‘ Aesthetics and History’ which is to be published in the next number of the College Art Journal, and one on Evelyn Sandberg-Vavala’s ‘‘ Uffizi Studies” published in the January 1949 Gazette des Beaux-Arts. Miss Elizabeth Mongan contributed six articles for the volume honoring Paul Sachs; an article for the Color Print Society on ‘Rockport,’ a colored lithograph by Stella Drabkin; descriptions of 27 illuminated miniatures to Pro- fessor Faye for the second edition of Seymour de Ricci’s “Census of Manuscripts in America.”’ An article on Rowlandson by David Keppel was published in the winter, 1949, number of The Art Quarterly. An article by James W. Lane entitled ‘Religious Art Exhibit’ appeared in the Interracial Review, and one on ‘‘Contemporary Religious Sculpture Exhibition” in the Catholic University Bulletin; he contributed two book reviews on ‘‘Van Eyck’s the Holy Lamb,” by Leo Van Puyvelde, and ‘‘Robert Louis Stevenson,’”’ by David Daiches, to the Catholic World, and one on “American Landscape Painting,” by Wolfgang Born, to the Magazine of Art. Charles M. Richards wrote a report on a code for intermuseum loans for the American Association of Museums. An illustrated catalog of the Gulbenkian Egyptian sculpture was issued for the opening of the exhibition, and Mr. Seymour prepared a pamphlet on the Michelangelo ‘‘ David,” which was placed on sale during its exhibition. The book of illustrations of the Mellon Collec- tion went to press in the late spring of 1949; work on the new National Gallery of Art catalog is at an advanced stage. The Publications Fund during the past fiscal year has continued to add new subjects to the supply of inexpensive color reproductions 36 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 offered to the public, including 11’’ x 14’’ color prints and color post cards. Five large collotype reproductions supplemented the already long list of subjects available. A silk-screen print of an anonymous fifteenth-century colored woodcut from the Rosenwald Collection was also published. The Gallery is continuing to meet the demand for illustrated catalogs of its various collections. The Mellon catalog is in process of publication, a third printing of the Kress catalog ordered, and a fifth edition of the Chester Dale catalog was published during the year. Two new publications were issued this year: an ‘Arts and Crafts Bibliography,” by Erwin O. Christensen, and a catalog of the ‘‘Egyp- tian Sculpture from the Gulbenkian Collection.” A group of engraved Christmas cards was added to the usual series of color and Rosenwald subjects. Final negotiations have been made for the printing in gravure of the book, ‘‘Masterpieces of Sculpture from the National Gallery of Art,” and it will be available by October 1949; the publisher now has the final manuscript for ‘‘ Made in America,”’ by Mr. Christensen; the Gallery received a stock of ‘Popular Art in the United States,”’ also by Mr. Christensen, which will go on sale on July 4, 1949; and “Pictures from America,”’ by John Walker, will shortly be published. EDUCATIONAL PROGRAM During the year approximately 15,000 persons attended the Gen- eral, Congressional, and Special Topic Tours, while over 20,000 attended the Picture of the Week. More than 18,000 came to hear the lectures and other programs in the auditorium. At least two- thirds of this lecture audience were regular attendants at these Sunday afternoon lectures. Many of them brought out-of-town visitors, and stated that this lecture series was becoming one of the Capital’s chief Sunday attractions. The motion picture, ‘The National Gallery of Art,” continues to be popular with clubs, educa- tional organizations, and similar groups. During the past 12 months, 19 persons borrowed this film. The publication of the monthly Calendar of Events, announcing Gallery activities, including notices of exhibitions, lectures, Gallery talks, tours, and concerts was continued during the year by the Educational Department. About 3,900 of the Calendar of Events are mailed each month. LIBRARY A total of 283 books, 221 pamphlets, and 31 periodicals were given to the Gallery; 494 books, 18 pamphlets, and 282 periodicals were purchased, and 40 subscriptions to periodicals were purchased. Exchanges with other institutions included 47 books, 114 pamphlets, SECRETARY'S REPORT 37 13 periodicals, and 420 bulletins. Of the 1,762 books borrowed and returned during the year, the Library of Congress lent 1,676 books to the Gallery on the usual interlibrary loan basis, and the remaining 86 books were borrowed from 25 public and university libraries. INDEX OF AMERICAN DESIGN During the year the Index of American Design continued to expand as the result of gifts and exchanges. Three hundred and thirty-six persons studied Index material at the Gallery; of this number, 301 were new users and 25 revisited the collection for study purposes. The use of photographs of Index drawings was increased by about 40 percent, with 1,796 photographs being sent out on loan, exchange, or purchase. Fifty exhibitions of original water-color renderings were circulated in 25 States. PRESIDENT TRUMAN’S INAUGURAL RECEPTION On January 20, 1949, the President’s Inaugural Reception was held in the National Gallery of Art. The Seventh Street ground floor and main floor lobbies were especially furnished and decorated for the occasion; the rotunda and the two garden courts were appro- priately decorated with flowers; under arrangements made by the White House staff, a platform was built in the West Sculpture Hall where the President addressed the guests who could not be received personally in the West Garden Court. Three sections of the Marine Band Orchestra played during the reception. The total number of guests was approximately 8,000. CUSTODY OF GERMAN PAINTINGS On April 6, 1949, the Gallery accepted custody of the 97 paintings from Berlin museums which had been on an exhibition tour of the United States, part of the group of 202 German paintings stored in the Gallery building by the Department of the Army from December 1945 to March 1948. After the last exhibition of this collection of paintings in Toledo, Ohio, the collection was brought to Washington and stored in the Gallery for about 2 weeks pending final shipping arrangements. On April 20, 1949, the collection was delivered to the Army for return to the American Zone in Germany. The exhibition of the Berlin paintings in 13 museums throughout the United States resulted in the collection of $303,605.35 through ad- mission fees and voluntary contributions for the relief of German children in the American Zone in Germany. ‘These funds were de- posited with the Gallery and were later disbursed in accordance with instructions received from the Department of the Army. During the tour 1,307,001 persons viewed the paintings, in addition to 964,970 38 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 who saw them during the time the paintings were on exhibition at the National Gallery of Art in Washington. CUSTODY OF GERMAN SILVER On January 7, 1949, the Gallery returned to the Department of the Army for transport to Germany the 44 sealed cases containing silver- ware and glassware and belonging to the Hohenzollern family. The cases had been stored in the Gallery since April 11, 1947. CUSTODY OF WHITE HOUSE FURNITURE On November 24, 1948, the Gallery accepted custody of certain items of paintings, sculpture, and furniture belonging to the White House for storage in the building until the repairs to the White House are completed. Shipments of these items started on December 3, 1948, and con- tinued for several days thereafter. At the present time there are 76 works of art—paintings and sculpture—stored in the Gallery’s storage rooms and 25 vanloads of furniture stored in the packing space on the main floor. The necessary arrangements for fire prevention, inspection, and fumigation have been established and are being carried out. NEW CONSTRUCTION During the past fiscal year, the Committee on the Building approved the construction in the southwest moat of a small workroom for the use of the gardening staff in maintaining and growing certain plants for the garden courts and landscaping. Later, when funds become available, it is planned to construct two small greenhouses adjacent to this workroom. The growth of the Gallery’s collections of works of art has been so rapid that all available exhibition space is now being utilized. As a matter of fact there are already several paintings which cannot be ex- hibited because there is no space in the present galleries. For this reason the Committee on the Building recommended that, to take care of the most urgent needs, the unfinished spaces 61-66 and 68-70, on the main floor, be completed as soon as funds are available. These galleries will be used for new acquisitions of paintings in the American and British schools and will also make possible some rearrangement in galleries already finished so as to make available additional space therein. The Committee on the Building also recommended that the so-called copyists’ room be finished to furnish office space for the Educational Department, which is now operating in rather cramped quarters. Funds have been generously made available from private sources to complete this work, and contracts have been entered into with Eggers SECRETARY'S REPORT 39 and Higgins, Architects, and Vermilya-Brown Company, General Contractors, for the completion of 12 galleries in these unfinished areas. The floor plan has been approved, and bids are now being taken from subcontractors. It is anticipated that actual construction will begin in August 1949 and that the work will be completed by May 1950. CARE AND MAINTENANCE OF THE BUILDING The usual routine work in connection with the care and maintenance of the building andits mechanical equipment was carried on throughout the year. The three older refrigeration compressors were completely dis- mantled and overhauled, including the purge compressors. Three chilled-water pumps, including the electric motors, were completely overhauled and realigned by the mechanical staff. Twelve supply fans were cleaned and repainted to protect them against corrosion, The structural steel base for the large 400-horsepower motor driving No. 2 Worthington refrigeration machine was strengthened in order that this large motor would remain in alignment. ‘To correct serious leaks in two of these machines, the technical staff successfully made and installed the necessary parts. The cornice metal lining at the top of the exterior wall of the building developed leaks, and approximately 50 percent of the joints in the metal lining were cleaned and soldered. In connection with the Inaugural Reception, the technical staff installed floodlights on three sides of the building, assisted the person- nel of the U.S. Army Signal Corps in the installation of a loud-speaker system on the main floor, and installed extra electric lines and water lines for the use of the caterer. The maintenance staff erected exten- sive checking facilities for the proper care of wraps. Twelve new display cases were constructed by the staff for the Gulbenkian Exhibition. Care and improvement of the Gallery grounds and other miscella- neous work progressed satisfactorily. Potted plants, totaling 2,366, which were used for decoration in the two garden courts, were grown in the southwest moat. In addition, over 350 large pots of chrysanthe- mums were also grown in this moat area, and these plants provided the decoration for the two garden courts during the months of October and November. COMMITTEE OF EXPERT EXAMINERS During the year the United States Civil Service Commission’s Committee of Expert Examiners, composed of staff members of the Gallery, aided in the drafting of standards for Civil Service positions in which a knowledge of the history of art is a basic requirement. The 866591—50-——4 40 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Committee also performed preliminary work in the preparation of the examination announcement for art positions which was dis- tributed by the Civil Service Commission with a closing date of April 19, 1949. From this examination registers of eligibles will be established for appointment to art positions in the Gallery and elsewhere in the Government. This will give the present incumbents, most of whom are serving indefinite war-time appointments, an opportunity to attain permanent status, and will also make available a greater number of qualified candidates. OTHER ACTIVITIES Forty-six Sunday evening concerts were given during the fiscal year, all concerts being held in the East Garden Court. A Mozart Festival of six concerts was given in the autumn with the highest attendance rate for the season. The five Sunday evenings in May were devoted to the Gallery’s annual American Music Festival. An estimated 50,000 persons attended these concerts. During the year the photographic laboratory of the Gallery made 17,709 prints, 1,342 black-and-white slides, 1,005 color slides, 3,873 negatives, in addition to infrared photographs, ultraviolet photographs, X-rays, and color separation negatives. A total of 3,500 copies of press releases, 128 special permits to copy paintings in the National Gallery of Art, and 117 special permits to photograph in the Gallery were issued during the year. OTHER GIFTS Gifts of books on art and related material were made to the Gallery library during the year by Paul Mellon and others. Gifts of money during the fiscal year 1949 were made by the Avalon Foundation and The A. W. Mellon Educational and Charitable Trust, and a cash bequest was received from the Estate of the late William Nelson Cromwell. AUDIT OF PRIVATE FUNDS OF THE GALLERY An audit of the private funds of the Gallery has been made for the fiscal year ended June 30, 1949, by Price Waterhouse & Co., public accountants, and the certificate of that company on its examination of the accounting records maintained for such funds will be forwarded to the Gallery. Respectfully submitted. Huntineton Cairns, Secretary. THe SECRETARY, Smithsonian Institution. APPENDIX 3 REPORT ON THE NATIONAL COLLECTION OF FINE ARTS Sir: I have the honor to submit the following report on the activi- ties of the National Collection of Fine Arts for the fiscal year ended June 30, 1949. THE SMITHSONIAN ART COMMISSION The twenty-sixth annual meeting of the Smithsonian Art Commis- sion was held in the Regents’ Room of the Smithsonian Building, on Tuesday, December 7, 1948. The members present were: Paul Manship, chairman; Alexander Wetmore, secretary (member, ex officio); George Hewitt Myers; George H. Edgell; Lloyd Goodrich; John Taylor Arms; Archibald G. Wenley, Gifford Beal, and Robert Woods Bliss. Thomas M. Beggs, Director of the National Collection of Fine Arts, and John E. Graf, Assistant Secretary of the Smithsonian Institution, were also present. The Commission recommended the reelection of Archibald G. Wen- ley, David E. Finley, Eugene E. Speicher, and Paul Manship for the usual 4-year period. The following officers were reelected for the ensuing year: Paul Man- ship, chairman; Robert Woods Bliss, vice chairman; and Dr. Alexander Wetmore, secretary. The following were reelected members of the executive committee for the ensuing year: David E. Finley, chairman, Robert Woods Bliss, and Gilmore D. Clarke. Paul Manship, as chairman of the Commis- sion, and Dr. Alexander Wetmore, as secretary of the Commission, are ex officio members of the executive committee. The Secretary summarized the status of exhibition and storage of the art objects of the National Collection of Fine Arts which at present are housed in space intended for the natural history collections in the Natural History Building. 33, 441. 68 ——————— 6, 092, 775. 69 otalsinvestInentsts 42 s—= ese aaeeee 2 ae eee 9, 141, 069. 65 146 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 CASH BALANCES, RECEIPTS, AND DISBURSEMENTS DURING FISCAL YEAR 19491 Cash balance/onphand! June’ 30) 194823225) 3 es See $563, 847. 37 Receipts other than Freer endowment: Income frompnvestMmentsas sae aes ee eee $156, 219. 10 Giftsyandtcontnbutions#s= 452s ose eee 48, 143. 71 Salesvor pubheatians= 2! ooe = (eee) es ewe ewes 33, 281. 09 IMS Cel Ane OUS) eRe ER Pe eek Cis | 11, 566. 10 Total receipts other than Freer endowment-_-__-___------- 249, 210. 00 Receipts from Freer endowment: Income fromanvestments== = 25a ae ee $282, 265. 48 Total receipts from Freer endowment-_-_-_-_-__--_--------- 282, 265. 48 TE GER eh es oe ee (Ee hie) Brie SR oa Oe 1, 095, 322. 85 Disbursements other than Freer endowment: AGministrations= 22 22 oe e eee eo eae ees $43, 422. 75 PST ORGLORIS eee ee I a erent Ee 45, 618. 12 Bio) 75 2 PRE eS Se. dee ee Se ee ees 3, 977. 10 Buildings—care, repairs, alterations__._______- 136. 00 @ustodiantieesNetos: Gees 2 28 ee 3, 293. 15 Miscellaneous 4222s. ano ees eta Sas 3, S22. 07 Mesearchesse. 22. Soe Une eee ae eee 127, 412. 84 Smithsonian Retirement System_____________- 3, 608. 28 Purchases of securities (net) _.__..-.-__-_____- 4, 508. 63 Total disbursements other than Freer endowment_-_-_-_-_-__- 235, 799. 64 Disbursements from Freer endowment: Salaries = 2 Areeet ee es eee ee $83, 480. 37 iRurchasestoncollectionsss.2 = — eee 125, 050. 00 Custodian fees etohaisa 2 2-2 eee 10, 858. 00 Miscellaneoud@oe ss soe ee cee oe a nee eee 26, 594. 80 Purchases of securities (net) __._._.._..________- 80, 631. 18 Total disbursements from Freer endowment__....__.-_--- 326, 614. 35 Investment of current funds in U. 8. Bonds________-__-__-.___- 2, 578. 13 fhotaldisbursements= = So o= 3226-2 sea ee eee 564, 992. 12 Cashibalante June30, 1049incteeabso cuss. sees ee eee 530, 330. 73 TOtalG: Ses oa55 he eee ee EE Fa en ee 1, 095, 322. 85 1 This statement does not include Government appropriations under the administrative charge of the Institution. SECRETARY’S REPORT ASSETS Cash: United States Treasury cur- Tent Account ae = see ae $360, 201. 95 In banks and on hand_----- 170, 128. 78 530, 330. 73 Less uninvested endowment fundshe 2s ea we eee 89, 826. 17 ED OE CE Travel and other advances___-_----------- 11, 585. 42 Cash invested (U.S. Treasury notes) ------ 502, 815. 37 Investments—at book value: Endowment funds: Freer Gallery of Art: Stocks and bonds__ $6, 059, 334. 01 Uninvested capital_ 33, 441. 68 ——_—_—__————_ 6, 092, 775. 69 Investments at book value other than Freer: Stocks and bonds_-__--- $1, 929, 118. 64 Real estate and mort- gage notes=—— 2 62, 790. 83 Uninvested capital____-_ 56, 384 49 Special deposit in U. S. Treasury. Interest at 6G; percent==ss====—"— 1, 000, 000. 00 —— 3, 048, 293. 96 147 $954, 905. 35 9, 141, 069. 65 10, 095, 975. 00 UNEXPENDED FUNDS AND ENDOWMENTS Unexpended funds: Income from Freer Gallery of Art endowment-_------------ Income from other endowments: Restricted == sees oa es ae ee $187, 425. 28 Goneral os besa ts ee Ok ao ee ee 85, 599. 74 Giltstindvpranite: 2. 5eie Sees io es ls ee Endowment funds: RreersGallery Of Att. o22s-. 2526s ee aon $6, 092, 775. 69 Other: Restrictedes sss) ee —— $1, 345, 776. 78 General 2 oe eek ee 1, 702, 517. 18 3, 048, 293. 96 393, 411. 62 273, 025. 02 288, 468. 71 954, 905. 35 9, 141, 069. 65 10, 095, 975. 00 148 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The practice of maintaining savings accounts in several of the Washington banks and trust companies has been continued during the past year, and interest on these deposits amounted to $824.74. In many instances, deposits are made in banks for convenience in collection of checks, etc., and later such funds are withdrawn and deposited in the United States Treasury. Disbursement of funds is made by check signed by the Secretary of the Institution and drawn on the United States Treasury. The foregoing report relates only to the private funds of the Institution. The Institution gratefully acknowledges gifts from the following: American Philosophical Society, for Iroquois research. W. W. Corcoran, for B. F. Starr. Eickemeyer Estate, for preservation, etc., of Rudolph Eickemeyer photographic collection. KE. P. Killip, for use of Department of Botany. National Academy of Sciences, for study of flora of Okinawa. National Geographic Society, expedition to western Panamé, Research Corporation. John A. Roebling, additional contribution for researches in radiation. The following appropriations were made by Congress for the Government bureaus under the administrative charge of the Smith- sonian Institution for the fiscal year 1949: Salariesiandvexpensessueseree i se eee oe Sao ae eee eee eae $2, 259, 000. 00 INStIOnalY SOOO sICO Ean K Shae sie eee SE Fs See 528, 848. 00 In addition, funds were transferred from other Departments of the Government for expenditure under direction of the Smithsonian Institution as follows: Cooperation with the American Republics (transfer from the State Weparvment). 28 es eee ewe nee eee ee meee eee ee eee $97, 960. 00 Working Fund, transferred from the National Park Service, Interior Department, for archeological investigations in River Basins throughout the United States_.._.........--.-.------- $118, 500. 00 The Institution also administers a trust fund for partial support of the Canal Zone Biological Area, located on Barro Colorado Island in the Canal Zone. The report of the audit of the Smithsonian private funds follows: SEPTEMBER 14, 1949. To THE Boarp or REGENTs, SMITHSONIAN INSTITUTION, Washington 25, D. C. We have examined the accounts of the Smithsonian Institution relative to its private endowment funds and gifts (but excluding the National Gallery of Art and other departments, bureaus, or operations administered by the Institution under Federal appropriations) for the year ended June 30, 1949. Our examina- tion was made in accordance with generally accepted auditing standards, and SECRETARY’S REPORT 149 accordingly included such tests of the accounting records and such other auditing procedures as we considered necessary in the circumstances. The Institution maintains its accounts on a cash basis and does not accrue income and expenses. Land, buildings, furniture, equipment, works of art, living and other specimens and certain sundry property are not included in the accounts of the Institution. In our opinion, the accompanying financial statements present fairly the position of the private funds and the cash and investments thereof of the Smith- sonian Institution at June 30, 1949 (excluding the National Gallery of Art and other departments, bureaus or operations administered by the Institution under Federal appropriations) and the cash receipts and disbursements for the year then ended, in conformity with generally accepted accounting principles applied on a basis consistent with that of the preceding year. Prat, Marwick, MircHe.y & Co. Respectfully submitted. Rozert V. FLEMING, VANNEVAR Bus, CLARENCE CANNON, Executive Commiitee. ae oxi oe ah rf rs be ay on me webrope sy a : - yah noes & Fa i" eater ee eT be a “ose Yon aor Twit 7 ans Se Oe | nd: eitedes " t ay ufstieet rt =e ei 9) alSerds ede ytups ater tical Firstly: ae WAY at idea: i Witgo easteiax nic més, Gwe lesieviloe ga seth ut = fas hea eiled S pene ous aul 9 ae i atc =) a = sad 52 | ll 5 I = e< Bei = ath VOUS droit) aluontaineg Lejanert,, gies tachment sill sipping 3 u - ory es ah ) >, 7 =f . 13} uo) To Toes itd dy MVE2 aS i r hile ifs 4 ty. Ds Ur Pitul of# ving 602 ce: ‘cn ni 7 Tee ow te ; J foros Ath, gnivnitss 9) Ube t e ny saint peliad iedliaaial aig wahAy (eR oaY i f Perkigiuiethe Siciliano i o> oo eae x z U: 8 ¢c o ee) mee: <—- er Pi) e > — Ficure 3.—Upper, our view of some spiral nebulae. The arrows indicate veloc- ities. Note that spiral B, which is twice as far from us as spiral A, is receding twice as fast. OC, three times as far, is receding three times as fast, and so on. Lower, the view of an observer on spiral B, considering himself to be at rest. It is the principle of relativity that he has just as much right as we do to con- sider himself at rest. He gets the same view as we do; all the spirals are receding from B with velocity proportional] to distance. Tracing the motions back in time (there is no evidence that the spirals are accelerating or decelerating) shows that all the spiral nebulae would have been near our galaxy between 2 and 3 billion years ago. The coincidence of this with the age of the earth and the age of the meteorites is too marked to need further comment—the whole universe seems to have started with a bang about 3 billion years ago! ‘ THE BEGINNING OF TIME This curious evidence that the spiral nebulae were all close to—if not entangled with—our galaxy 8 billion years ago, means that the formation of the solar system at that time probably took place under conditions somewhat different from those of today. To be sure of the reasoning, we must examine the conditions of 3 billion years ago more carefully; it was this reexamination which led, in 1945, to the most bizarre suggestion of all in this field already rich in speculation. It was put forward by the English biologist, J. B. S. Haldane, and is based on a new theory—or philosophy—of relativity proposed in 1932 by the English mathematician, E. A. Milne. First we shall speak of Milne and his brand of relativity. To make the reasoning clear we must start with Einstein’s earlier relativity theory which links space and time in such a way that if one observer is moving at constant velocity past another his measurements of distances and time intervals will differ from those 172 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 of the first observer, although the relation of time and distance is such that they both observe the same laws of physics. LHinstein formulated his relativity on the philosophy that it is simply impos- sible to tell which observer is ‘‘at rest.’”? Complicated as it sounds, this scheme has been developed to form a logically complete theory in terms of mathematical transformations. Milne extended the established principles of relativity in his “cosmological principle,”’ which is, in effect, an assumption that the view of the whole universe from one spiral nebula must be the same as the view from any other. Moreover, he has redefined distance measurements in terms of the travel time of light signals, as in radar ranging, thus reducing both time and distance measurements to readings of clocks, in principle. ¢ e e ‘ ¢ . ’ ¢ eo e ¢ e oe ° e e e e o e ¢ 7 e a e e e o . . ¢ e e a e ¢ eo ¢ e e ‘ ¢ ¢ o ° e ’ ° eo eo e ¢ Us o e e ° 2 , e ¢ ‘ eo 6 oe eo Figure 4.—Milne’s picture of the universe. If all measurements are made in atomic time, on Milne’s theory, the universe started expanding from a point 3 billion atomic years ago. As we see it now the spiral nebulae shown in the left diagram are all moving away from us and (if we could see far enough) would be much more numerous near the “edge.”” At this edge the velocity of recession is equal to the velocity of light, so we can never hope to see the edge itself. On the other hand, if clock time is used for all our measurements, the universe is static and the spiral nebulae, as shown on the right above, are uniformly distributed on to infinity. The more distant nebulae are redder be- cause we see them as they were many years ago with “‘slow” atoms. The “edge’’ of this picture comes when this reddening gets so extreme that galaxies are no longer visible. Milne then raises the disturbing question: How are we sure that our clocks are reading constant intervals of time? In fact, the slow- ing down of the earth’s rotation (which is normally our ‘master clock’’) has been measured as one-thousandth of a second per century by comparison with the planets, and we have no philosophically sound assurance that the planets keep “‘perfect time.” The cosmological principle leads mathematically to two kinds of time, one of which is speeding up relative to the other. Milne has shown that pendulum clocks, the earth, and the planets keep ‘‘dy- ORIGIN OF THE EARTH—PAGE tzZ3 namic’”’ or clock time, while vibrating atoms and radioactive decay have constant period only in “kinematic”’ or atomic time. There is no philosophical reason for choosing one kind as the “correct” time; if we used a pendulum clock to time atoms we would find, after a very long interval, that the atoms are gradually speeding up in their vibration, if we used an atomic clock we would similarly find that the planets are slowing down in their orbits. If this is correct—and no one has yet proved it otherwise—the age of the earth is 3 billion atomic years as determined from radioactive decay, but it is many more clock years, since in the past the clock year was shorter than the atomic year. (They are equal at present— by definition.) The coincidence between the age of the earth and the time of reces- sion of the spiral nebulae Milne explains as a result of the difference in these two kinds of time. Since the light we observe from a spiral 100 million light-years away left there 100 million years ago, we are seeing the atoms there ticking off the units of atomic time in use 100 million years ago. Compared to our present atoms, these early atoms ran slow; as a result, the light they emitted is redder than the light emitted now by similar atoms on the earth. From this effect and his cosmological principle, Milne calculates that in the past infinite number of clock years there were 3 billion atomic years. The origin of the earth, and the time when all the spirals were close to our galaxy, both of them 3 billion atomic years ago, therefore occurred at the beginning of time (since one could hardly expect more than infinite time on the clock scale). Now for Haldane’s suggestion, which he calls ‘A Quantum Theory of the Origin of the Solar System.” It is based, as its name implies, on the well-established quantum theory of radiation, and on a mathe- matical result of Milne’s theory: that the universe, as measured in atomic time, has expanded with the velocity of light, starting from a point of zero radius 3 billion atomic years ago. Since the universe started from zero radius, Haldane was able to pick an early enough instant, just a fraction of a second after the start of atomic time, when the whole universe was but a fraction of an inch in diameter—much smaller than the wave length of visible light— smaller, by far, than the wave length of X-rays or gamma rays. (These fractions are too small to write out easily; the first requires 72 zeros after the decimal point, the second, 62!) ‘The wave lengths of radiation in existence in this small universe could scarcely have been bigger than the universe itself, Haldane reasoned, therefore the only radiation in existence was of these incredibly short wave lengths. But the basic principle of the quantum theory is that radiant energy comes only in packets, or ‘quanta,’ inversely proportional to the wave length in size. So, at this early instant all radiation was in 174 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 giant quanta of very small waves. And the energy of one of these giant quanta can easily be calculated as sufficient to knock one or more planets out of the sun. The even smaller waves at a somewhat earlier instant would have been in quanta with sufficient energy to tear apart stars, and even earlier, to tear apart the galaxies from some primordial globe of matter. The details of this remarkable suggestion have been carried no farther, but Haldane’s investigation points up one important general fact: whether or not Milne’s new relativity is accepted, conditions at the time of the origin of the solar system were probably consider- ably different from those today. If Milne’s cosmology is accepted, the relationship between radiation and matter was most radically different. It may seem that this iast and most fantastic speculation— which can neither be completely explained nor fully evaluated here— contradicts our former conclusion that the solar system was formed from a rotating nebula of gas and dust. However, the condensation of the planets and the distribution of angular momentum (which have been so difficult to explain in all previous theories) may follow from further mathematical investigation of the first second of atomic time. In fact, if the details can be worked out rigorously, Haldane’s suggestion may lead to confirmation of Milne’s cosmology, which is as yet lacking. In an echo of the introductory remarks it scarcely needs to be emphasized that we have no complete theory of the origin of the earth. The reader may be impressed with the diverse investigations involved and with the promise of the latest speculations; or he may notice the infinite regression implicit in any question of origins: if the planets were formed from dust or planetesimals, whence came the dust or planetesimals? if the dust and planetesimals came from a primordial nebula, whence came the primordial nebula? if the primor- dial nebula was formed by the absorption of a giant quantum by a fragment of matter, whence came the original matter and radiation in the universe? and so on, ad infinitum (clock time). THE 200-INCH HALE TELESCOPE AND SOME PROBLEMS IT MAY SOLVE! By Epwin Hussite Mount Wilson Observatory [With 10 plates] In 1609 Galileo turned his telescopes toward the sky. His favor- ite—it was the fifth, finished within 6 months of the first trial—was about 5 feet long and had a lens about 2 inches in diameter. It magnified nearly 30 times and its light-gathering power was equal to about 80 human eyes. He called it ‘Old Discoverer,” and with it he saw mountains on the moon, phases of Venus, four moons of Jupiter, and stars innumerable beyond the limit of the unaided eye. It was then that the explorations of space began—the explorations that have swept outward in wave after wave as telescopes developed, until in our time we study a region of space so vast that it may be a fair sample of the universe itself. Today there is nearing completion a new telescope, far more powerful than any previously made, and it is proper to consider its significance both as an engineering achieve- ment, and as an instrument for further explorations. With this end in view, I propose to discuss briefly the development of telescopes in general, the 200-inch in particular, and some of the problems it may help us to solve. Galileo’s optic tubes with single-lens objectives grew rapidly into telescopes from 20 to 25 feet long with lenses 2 to 3 inches in diameter. There the development stopped, for practical purposes, because of the engineering difficulties with still longer tubes. The longer focal lengths were considered desirable in order to over- come color difficulties. With a single lens, each different color was brought to a focus at a different distance from the lens. Hence the image, when focused for any particular color, was blurred by the out- of-focus images in other colors. The long telescopes represented an attempt to spread out the images of different colors over so long a distance that one color could be focused with minimum interference 1 Alexander F. Morrison lecture, delivered in Pasadena, Calif., April 8, 1947. Reprinted by permission, with slight alterations, from Publications of the Astronomical Society of the Pacific, vol. 59, No. 349, August 1947. 175 176 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 from the others. Telescopes 100, 150, and even 200 feet long were actually constructed, with lenses from 3 to 6 inches in diameter. These monstrous instruments, however, were too unwieldy for use, and the real work during the first century and a half after Galileo’s time was done with the smaller telescopes. Finally, in the middle of the eighteenth century, the color problem was solved by replacing the single-lens objective with a compound objective, each of whose separate components, made of different kinds of glass, canceled out most of the color effects of the other. These color-free (achromatic) lenses gave much better images and permitted the use of relatively short tubes for a lens of a given diameter. Tele- scopes immediately entered a new period of growth which culminated in the 40-inch lens, with a focal length of 63 feet, at the Yerkes Observatory in Wisconsin. The 40-inch was finished in 1892, and since that time developments have concerned lenses for special pur- poses rather than for greater light-gathering power. For technical reasons, it seems unlikely that larger lenses will be made in the foreseeable future. This greatest of all lenses had been ordered originally by a group of enthusiasts here in southern California in connection with a plan for a ‘University System.” The project did not fully materialize, and the unfinished telescope was bought and completed by the University of Chicago. The very large telescopes of recent times, in which light-gathering power is the most important consideration, are all reflectors, not refractors. The light is funneled to a focus, not by refraction through a convex lens but by reflection from a concave mirror. ‘These tele- scopes are free from color effects because all colors are reflected in the same way. The first reflector was made by Isaac Newton in 1672, in a deliberate effort to avoid the color troubles of single-lens refractors. His first model had a burnished metal mirror, about an inch in diameter, figured to a concave spherical surface, and mounted at the bottom of a tube about 6 inches long. The image, which would lie in the middle of the upper end of the tube, was thrown to the side by a small plane mirror set at 45° to the axis, just below the focus. Newton presented the toy to the Royal Society, where it may still be seen, sitting on a volume of his famous Principia. Although Newton’s reflector avoided the color problem, it suffered another defect, known as spherical aberration, arising from the spherical surface of the main mirror. It was not until 50 years later when Hadley, in 1722, found a method of parabolizing concave mirrors, that the development of reflectors finally got under way. About 90 years ago metal mirrors were replaced by glass, silvered on the front surfaces. In our time aluminum has been substituted for silver, 200-INCH HALE TELESCOPE—HUBBLE ZAC low-expansion glasses have been developed, methods of parabolizing have been perfected, and engineering problems of constructing telescopes have been solved as they arose. The 40-inch refractor was installed at Yerkes under the direction of George E. Hale. He clearly saw that, regardless of the success of this telescope, the quest for still greater light-gathering power depended upon mirrors rather than lenses. Refractors were pref- erable for certain types of work (including, for instance, visual resolution of double stars, precise measurement of position, wide- angle photography, ectc.), but for light-gathering power, with all that it implies, the future lay with the reflector. Because the reflections are from the front surfaces, transparency and absolute homogeneity of the glass are not demanded; the mirror may be supported from the back and sides, instead of from the rims alone as in the case of lenses, and, of course, there are no color effects. Hale took the lead in America in encouraging the development of large reflectors. A 24-inch of unusual perfection was made by G. W. Ritchey and installed at Yerkes. It proved so successful that plans for a 60-inch were immediately set in motion. When Hale left Yerkes to establish the Mount Wilson Observatory, he was able to transfer Ritchey and the unfinished 60-inch mirror to Pasadena, where the telescope was completed in 1908. The work of the 60-inch on Mount Wilson so fully justified the faith in larger reflectors that plans for a new one were immediately made, this time for a 100-inch mirror. This reflector, completed during the first World War, marked an important epoch in the history of astronomy. It is still the greatest telescope in operation. Four large reflectors with mirrors from 60 to 84 inches have since been completed (two in Canada and two in the United States), and others, including a 120-inch for Lick Observatory, are in process of planning or construction. The 100-inch opened up new fields of investigation of the very first importance, and furnished glimpses of even richer fields beyond. If more light-gathering power were available, these more distant fields could be explored. In the face of this challenge the possibility of larger telescopes was the favorite topic of conversation among astron- omers at Mount Wilson, and presumably at other places as well. We talked of 200 inches, or 300, and even dreamed of still more light. One of the group, F. G. Pease, drew tentative designs for a 300-inch, and demonstrated that the engineering features were not impossible. Again Hale took the lead. Through his efforts funds were secured in 1928 in the form of a gift from the International Education Board to the California Institute of Technology for the establishment of an astronomical observatory and laboratory. An Observatory Council, with Hale as chairman, and with the greatest experts in the country as advisers, administered the details of the project. When Hale 178 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 dropped out, Max Mason took over the chairmanship. It was decided that a 200-inch reflector was as bold a step beyond the 100-inch as could be justified in view of the unknown problems, both optical and engineering, that might be encountered. Laboratories and shops were erected on the California Institute campus, a site for the ob servatory was found on Mount Palomar, a disk of pyrex glass was achieved by the Corning Glass Company, and the project proceeded steadily until it was interrupted by the war. Work was resumed soon after VJ-day, and the telescope has since been completed. The proper fields for the 200-inch are determined primarily by its immense light-gathering power. Because adequate consideration of all the possible applications would require more time than is now available, I propose to limit the following discussion to three typical problems. These problems are, first, the existence of canals on Mars; second, the relative abundance of the chemical elements in stars; and, third, the large-scale structure of the universe. Each problem represents a particular aspect of light-gathering power, namely, resolution, dispersion, and depth penetration. As a brief introduction, let me comment on the telescope itself. The mirror intercepts a beam of light 200 inches or 17% feet in diameter—in other words, it gathers as much light as a million human eyes, or four 100-inch reflectors. It funnels this light to an image at the primary focus, 55% feet in front of the mirror. There an image of the sky is formed such as you may see on the ground glass of a camera. This image may be examined visually with a microscope, recorded on a photographic plate, analyzed with a spectrograph, or studied by other techniques. Actually, most of the work will consist of direct photography or spectrum analysis. By using long time exposures, it is possible to photograph stars or nebulae several times fainter than can be seen in the eyepiece. For this reason, the 200- inch is best described as a huge camera. Now let us consider some typical problems for the 200-inch. I shall start with a problem concerning a member of the solar system. The telescope will not be turned on the sun because of temperature effects— in some ways it would act as a burning glass. Nor does it offer any unique advantages for the study of the moon. In that field it will serve merely to improve data of a kind that can be got nearly as well with several other telescopes. In the field of planetary photography, however, the opportunities are unique because, for the first time, it may be possible to photograph all that the eye can see with a telescope of moderate size. An immediate application is to the highly contro- versial question of canals on Mars. The canals are described as very fine, dark lines running along great circles, sometimes doubled, and often converging or crossing at spots called “‘oases.”’ Such fine, hairlike patterns, superposed on the back- 200-INCH HALE TELESCOPE—HUBBLE 179 eround of large, well-known markings, have been recorded by various trained visual observers, using telescopes of all sizes from 6 inches up- ward, during the whole of the past 70 years since Schiaparelli first reported them. ‘The canal systems, if real, would almost necessarily imply the existence now or in the past of intelligent beings on Mars. But other trained observers, again using telescopes of all sizes, report no trace of canals. KE. E. Barnard, perhaps the greatest of the American visual observers, studied the planet over many years with the then largest telescopes in the world, including the 36-inch refractor at Lick, the 40-inch at Yerkes, and the 60-inch reflector at Mount Wilson, and, although he saw an immense amount of detail, he found no canals. The two groups of observers flatly contradict each other, and since the observations are personal impressions neither group can demon- strate the validity of its assertions. Evidently the controversy must be resolved by photography. Once photographs are available on which the canals should appear if they are real features of the planet, the question will be settled beyond reasonable doubt. The test has not been possible as yet because existing equipment, although closely ap- proaching the required standards, does not fulfill them. The 200- inch, however, should meet all the necessary conditions and settle the question. The problem is as follows: Mars is a small object. The image at the primary focus of the 100-inch is less than 5 inch at the most favorable oppositions, and less than % inch at the long Cassegrain focus. In order to get an image large enough to serve as a critical test of fine markings (i. e., to make the resolution of the photographic plate comparable with the optical resolution of the telescope) it is necessary to use an enlarging lens. Thus the total light collected by the telescope is spread over a much larger, and correspondingly fainter, image. Furthermore, because of the atmosphere on Mars, it is neces- sary to photograph the surface markings through deep orange or red filters, still further reducing the effective brightness of the image. The reduction is so great that photographs with existing telescopes require time exposures instead of snapshots. The exposures may be only a second (or even a fraction of a second, with the 100-inch) but they are long enough to permit the dancing or shimmering of the image to smear out the finest detail. You doubtless realize that a telescope magnifies the twinkling of stars along with everything else. Critical observations are restricted to periods of maximum steadiness (good seeing, as it is called), and even then, the shimmer is appreciable under high magnification. The eye can “hold”? an image under these conditions, but photography with time exposures is helpless. The shimmer smears out the fine details. It is for this reason that, in the case of fine markings such 180 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 as the canals on Mars are said to be, the eye can see more than the photographic plate can record. However, the 200-inch will collect so much light that, for the first time, it will be possible to photograph enlarged images through filters with snapshots. These exposures will be short enough to catch a dancing image at the end of a flicker—when it is momentarily at rest as it reverses direction. If many exposures are snapped on a movie film, a certain percentage of them may be expected to record what the eye can see (at least with telescopes of moderate sizes). There is much more to the story, but it is too technical for the present discussion. But it can be said with some confidence that the 200-inch may settle the long-standing controversy concerning the canals on Mars. The second problem I have selected for discussion involves spectrum analysis. You know, of course, that light reaches us as a jumble of waves of all different wave lengths, each representing a different color. It is possible, with prisms or gratings, to spread the colors out into an ordered sequence or spectrum, running from the long waves of thered to the short waves of the violet, and beyond in either direction. Such spectra of stars and nebulae show phenomena of profound significance at certain particular wave lengths. Spectrum analysis involves the isolation and study of these particular regions. Your radio offers an analogy. With the tuning dial you run along the spectrum of radio waves and isolate a particular wave length in order to hear a particular station which is broadcasting on that wave length. If there were no tuning device, and you heard all programs at once with a nonselective receiver, the result would be bedlam. The step from such a nightmare to the clear reception of messages from indi- vidual stations suggests the nature of the step in astronomy from the study of integrated light to spectrum analysis. Light from stars and nebulae originates in atoms. There are as many kinds of atoms as there are chemical elements, and the atoms may have various stable states. Each stable state of each kind of atom represents a set of broadcasting stations, sending messages concerning the nature of the atoms and the physical conditions under which they exist. By tuning in and reading these messages, it is pos- sible to identify chemical elements, to determine temperatures, pres- sures, and other physical attributes, and even to measure motion in the line of sight (radial velocity). But in order to read the messages clearly it is necessary to achieve precise tuning—that is, to spread out the spectrum on the maximum possible scale. It is here that the great light-gathering power of the 200-inch offers new possibilities. 200-INCH HALE TELESCOPE—HUBBLE 181 The length over which a spectrum can be spread, and still remain bright enough to be photographed, depends upon the brightness of the object. The sun has been spread out over a spectrum about 50 feet long from red to violet; the brightest stars, over about 3 feet, and the faintest naked-eye stars over about 1 foot. The shortest spectra giving useful information are about one-tenth of an inch long, and have been obtained from stars and nebulae about a hundred thousand times fainter than the faintest naked-eye stars. With the 200-inch, all the stellar and nebular spectra can be lengthened about four times, and consequently the analysis can be carried out much more precisely than was hitherto possible. One new field, now faintly glimpsed, can be explored rather fully. The important data are the relative abundances of the different chem- ical elements in different kinds of stars. These data are derived from the comparative study of the different stations (or lines) due to differ- ent chemical elements in a spectrum, and require the longest practical spectra (the highest possible dispersion) for adequate analysis. There is reason to believe that more than 99 percent of the atoms in the universe are hydrogen. Even by weight, hydrogen, with the simplest and lightest of all atoms, probably contributes a large frac- tion of the total matter in the universe. There are insistent sugges- tions that the relative abundance of hydrogen varies considerably from star to star. There is also some reason to suppose that the rela- tive abundance of other elements does not vary widely in the stars, although the physical conditions of the stars do vary widely (from giants to dwarfs, from hot blue stars to cool red stars). The supposi- tion rests mainly on negative evidence and requires further study with powerful instruments. It is believed that the 200-inch alone can adequately explore this field, now dimly outlined with existing telescopes. What is now sug- gested by analysis of three or four of the very brightest stars can be critically tested in these objects, and the study can be extended in a comparable way over a large sample collection of stars in general. We cannot predict the final results of the exploration. They may represent the next major chapter in the development of our knowledge of the universe, or they may prove to be relatively trivial. But the unexplored field looms as a challenge, and the challenge will be met. The data are immensely important because they bear directly on two very fundamental problems, namely, the source of stellar energy and the origin of chemical elements. Geologists, studying the history of the earth’s surface, assure us that the sun has been pouring out energy at a fairly constant rate over the last several hundred million years at least. Possible sources for the unfailing supply were not only unknown but were unimagined, 182 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 until the modern science of nuclear physics was developed. Now ex- planations may be sought over a wide range of nuclear reactions giv- ing various lifetimes to stars up to the limit set by Einstein’s famous formula, energy =mass X (velocity of light)? For instance, if the whole of the sun’s mass were transformed directly into energy, the sun could radiate at the present rate for a million million years. But, if nuclear reactions supplied the energy, the possible lifetime (with the present rate of radiation) would be reduced according to the particular reaction involved. The nuclear reactions, in general, produce the transmutation of elements—the old dream of the alchemists. The most plausible of the current theories concerning the source of the sun’s energy, pro- posed by H. Bethe, is based on the carbon cycle in which, because of the presence of carbon at temperatures found in the sun, hydrogen nuclei may combine to form helium, releasing energy in the process. One test of the theory is furnished by a comparison of the relative abundance of the different isotopes of carbon actually observed in the sun with the relative abundances involved in the carbon cycle. In a vaguely analogous way, it is possible to speculate on the build- ing up of all elements from the primitive hydrogen atoms, and these speculations may be guided by the observed relative abundances of elements in stars of widely different physical characteristics. Thus the data on abundances, derived from large-scale spectra, bear directly on all theories concerning the source of stellar energy, the origin of chemical elements, the past history of the universe, and its future evolution. The third unique field of investigation for the 200-inch is cos- mology—the structure and behavior of the universe as a whole. As- tronomers hope that the observable region of space—the region that can be observed with telescopes—is a fair sample of the universe, and they attempt to infer the nature of the universe from the observed characteristics of the sample. The 200-inch, because of its great light-gathering power, should penetrate into space about twice as far as the 100-inch, and consequently will permit us to explore a volume of space about eight times that now available. The probability that the observable region may be a fair sample of the universe will thus be greatly increased. It was the 100-inch that opened this new field and prepared the way for the new telescope. The picture developed rather suddenly during the 1920’s. The sun with its family of planets seems isolated and lonely in space, but we know that it is merely one of the stars— one of several thousand million stars which, together, form the stellar system. This system is a swarm of stars which drifts through space Smithsonian Report, 1949.—Hubble PLATE 1 DOME OF THE 200-INCH TELESCOPE ON PALOMAR MOUNTAIN Smithsonian Report, 1949.—Hubble PEATE 2 200-INCH HALE TELESCOPE Meeting of the American Astronomical Society and the Astronomical Society of the Pacific on July 1, 1948. Smithsonian Report, 1949.—Hubble PLATE 3 MARS Upper: Photograph with 100-inch reflector, September 2, 1924. Lower: Drawing by Pettit with 20-inch reflector, July 12, 1939. These typical pictures illustrate the fact that as vet photography does not furnish an objective test of the existence of “canals” on Mars. Smithsonian Report, 1949.—Hubble EXTRAGALACTIC NEBULAE These nebulae are examples of the stellar systems which serve as landmarks in the exploration of the universe. The group above (NGC 3185, 3187, 3190, 3193) is at a distance of about 8 million light-years and appears to be receding from us at the rate of about 850 miles per second. Smithsonian Report, 1949.—Hubble PLATE 5 NGC 2261 The first photograph made with the 200-inch Hale telescope. ZG VWaHYV GSLOATAS NI NOIS3Y 9 ALVI1d 2[999H— 6r6| ‘340dey UeluOsy{IUIG Smithsonian Rerort, | PLATE 7 MESSIER 87 (NGC 4486) Smithsonian Report, 1949.—Hubble PLATE 8 NGC 5204 Smithsonian Report, 1949.—Hubble PLATE 9 NGC 3359 Smithsonian Report, 1949.—Hubble PEATE 0 MESSIER 3 (NGC 5272) 200-INCH HALE TELESCOPE—HUBBLE 183 as a swarm of bees drifts through the air. From our position some- where within the system, we look out through the swarm of stars, past the boundaries, into the universe beyond. Those outer regions are empty for the most part—vast stretches of empty space. But here and there, scattered at immense intervals, we find other stellar systems comparable with our own. They are so remote that individual stars can be seen only in a few of the nearest systems. In general they appear as faint patches of light, resembling tiny clouds, and have long been called by the Latin word for clouds— that is, ‘‘nebulae.”’ We now know that these nebulae are huge stellar systems averaging about a hundred million times as bright as the sun. They are the true inhabitants of space—vast beacons that serve as landmarks for the exploration of the universe. We see a few that appear large and bright. These are the nearer nebulae. Then we find them smaller and fainter in constantly increasing numbers, and we know we are reaching out into space farther and ever farther, until, with the faintest nebulae that can be detected with the largest telescope, we have reached the frontiers of the observable region. This region has been explored with the 100-inch out to distances so remote that light, speeding at 186,000 miles per second, requires 500 million years to make the journey. Thus the observable region at present is a sphere, centered on the observer, with a radius of about 500 million light-years. Throughout this sphere about a hundred million nebulae are scattered, each a stellar system comparable to our own system of the Milky Way. The study of this observable region as a sample of the universe has led to the recognition of two large-scale features. The first feature is homogeneity. The nebulae are scattered singly, in groups, and even in great clusters, but when very large volumes of space are com- pared, their contents are found to be quite similar. On the grand scale, the observable region appears to be very much the same, in all directions and at all distances. The second characteristic is the fact that light waves from distant nebulae seem to grow longer in proportion to the distance they have traveled. It is as though the stations on your radio dial were all shifted toward the longer wave lengths in proportion to the distances of the stations. In the nebular spectra the stations (or lines) are shifted toward the red, and these red-shifts vary directly with dis- tance—an approximately linear relation. The red-shifts are most easily interpreted as evidence of motion in the line of sight away from the earth—as evidence that the nebulae in all directions are rushing away from us, and that the farther away they are, the faster they are receding. This interpretation lends itself directly to theories of an expanding universe. The interpretation is 866591—50——13 184 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 not universally accepted, but even the most cautious of us admit that red-shifts are evidence either of an expanding universe or of some hitherto unknown principle of nature. The two observed characteristics of the observable region, namely, the approximately uniform distribution and the approximately linear law of red-shifts, must be satisfied by any theory of the universe. They are the only observational results on the grand scale that can be used as tests. They serve to eliminate many theories formerly developed on insufficient data, but several modern theories survive the tests. These latter theories all permit the observed features in a limited region near the observer but they predict that departures from the simple approximate laws of distribution and of red-shifts will be found when the measures are extended to greater distances. These departures differ from theory to theory, and, if the measures can be extended to the necessary distance, will distinguish the correct theory from the false. Thus the most important observational problems in cosmology may be described as the small, second-order effects of great distances. The nebulae appear to be distributed in a roughly uniform manner and the red-shifts appear to be roughly proportional to distance, out to the limits of the 100-inch. The next step is to determine these features more precisely over the limited range of the 100-inch and approximately out to far greater distances. Attempts have been made to attain the necessary precision with the 100-inch, and the results appear to be significant. If they are valid, it seems likely that red-shifts may not be due to an expanding universe, and much of the current speculation on the structure of the universe may require re-examination. The significant data, however, were necessarily obtained at the very limit of a single instrument, and there were no possible means of checking the results by inde- pendent evidence. Therefore the results must be accepted for the present as suggestive rather than definitive. The problem is essentially one for the 200-inch. This new telescope will penetrate into space out to a thousand million light-years, and the second-order effects of great distance will be so conspicuous that they cannot be missed. As a particular and final example, let me mention the effects of increasing red-shifts on apparent brightness. It is well known that a rapidly receding light appears fainter than a similar, but stationary, light at the same momentary distance. The reason is that the stream of light-quanta from the moving light is thinned out by the recession so that fewer quanta per second reach the observer. Since brightness is measured by the rate of arrival of quanta, the receding light appears abnormally faint. 200-INCH HALE TELESCOPE—HUBBLE 185 Actually the dimming factor (the reduction of apparent brightness) is a simple fraction represented by velocity of recession divided by the velocity of light. Recession at one one-hundredth the velocity of light reduces the apparent brightness by 1 percent; at one-tenth the velocity of light, by 10 percent, and so on. Thus the effects of recession would be negligible until velocities of several hundred miles per second were reached. The effects would be appreciable at a few thousand miles per second, and conspicuous at several tens of thousands of miles per second. If red-shifts are evidence of actual recession, the dimming factors should become appreciable near the limits of measurement with the 100-inch and should be conspicuous near the limit of the 200-inch. At the very limits of direct photographs with the 200-inch, the factor should approach the order of 40 to 50 percent, and should be unmis- takable. We may predict with confidence that the 200-inch will tell us whether the red-shifts must be accepted as evidence of a rapidly expanding universe, or attributed to some new principle of nature. Whatever the answer may be, the result will be welcomed as another major contribution to the exploration of the universe. I have mentioned the three specific problems of canals on Mars, relative abundance of chemical elements in stars, and the nature of the red-shift, because they illustrate the unique powers of the 200- inch telescope in three aspects, namely, resolution, dispersion, and space penetration. Because these problems are of first importance, and can be solved, they, together with others of a similar kind, will be included in the initial research programs. The solutions of these problems alone will fully justify the construction of the telescope. But such a program is merely a logical beginning—the first carefully considered stage in the exploration of vast unknown regions of the universe. As the darkness is pushed back, greater problems will doubtless emerge which we cannot now foresee. FIRST PHOTOGRAPHS WITH THE 200-INCH HALE TELESCOPE? The first photographs with the 200-inch Hale reflector on Palomar, made under normal observing conditions, confirm the most optimistic predictions of its designers. Such a statement, as usual, requires some explanation. The photographs were made as routine tests to record progress in the tedious program of adjustments. Seeing was never better than “‘average,”’ the aluminum coat was dusty and grimy, and the mirror showed a turned-up edge. These handicaps, of course, 2 A later article by Dr. Hubble from the Publications of the Astronomical Society of the Pacific, vol. 61, No. 360, June 1949, 186 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 will be eliminated or avoided in time, but during the tests they caused some loss of light and appreciable loss of definition. Nevertheless, the test plates record stars and nebulae fully 1.5 magnitudes fainter than the extreme limit of the 100-inch reflector on Mount Wilson. The faintest star images, on the better plates, were, however, a little more than 1 inch in diameter, and, at the threshold, it was sometimes difficult to distinguish with certainty between stars and nebulae. Thus the 200-inch has registered already the full gain in light- gathering power corresponding to the size of the main mirror. The slight additional gain that may be expected with a clean, sensibly perfect mirror surface will be accounted for by the absence of a Newtonian flat and by the very transparent sky over Palomar. The greatest improvement in the future will be in definition, as indicated by the size of faint star images. Definition is very sensitive to the seeing, and, while the test plates approached the definition to be expected under average conditions, they indicated that the mirror is not yet in shape to operate at maximum efficiency on the rare nights of fine seeing. The trouble arises from the turned-up edge and can be eliminated by the retouching now in progress. The improved definition will be significant, particularly for distinguishing nebulae from stars at the threshold of long exposures. In the higher latitudes, the telescope records many more nebulae than stars. The turned-up edge was well known from Hartman tests, and its effects could be predicted with some confidence. The photographs were made primarily to confirm and record these effects. However, the first plates were so impressive that a set of full exposures was made to serve as a record of performance before the mirror was removed for retouching. About 60 photographs were assembled over the 3 months from January 26 to April 28, as opportunities arose during the normal program of adjustments. Of this number, perhaps half a dozen represent full exposures under average seeing conditions, and a like number show good performance with reduced apertures or with a Ross correcting lens for enlarging the usable field. Selections from the files are illustrated in plates 5 to 10, and comments on them are given below. The full aperture (200-inch) and Eastman 103a—O emulsions were used in all exposures except those to which special references are made in the comments. The scale of the original negatives is about 1 mm=12’’. Plate 5, NGC2261; R.A.=6"36"6, Decl. =+8°47’ (1950); Jan. 26, 1949; 15 min. exposure, poor seeing; enlarged 34x. This plate shows the first of the photographs with the Hale tele- scope. It is recorded as PH—-1-H (i. e., Palomar, Hale, No. 1, followed by the initial of the observer). It was made under poor conditions as a preliminary test of the mechanical operations and procedures involved in direct photography at the prime focus. The trial was 200-INCH HALE TELESCOPE—HUBBLE 187 successful, except for the large size of the star images produced by the poor seeing. The exposure was made on January 26, 1949, about 10 p. m., after waiting more than a week for a break in the weather. The object, NGC2261, is a well-known, variable galactic nebula—a, comet-shaped mass with the variable star, R Monocerotis, at the apex. Plate 6, S.A. 57; R.A.=13"6"3, Decl.=-+29°37’ (1950); Jan. 27, 1949; 60 min. exposure, seeing average; enlarged 7% ; center is 2/5 N. and 3/4 W. of BD+30°2371. This selected area contains one of the most reliable magnitude sequences available for faint stars extending to the 21st magnitude. Exposures of 1 minute registered stars to about 19.7, and of 3 minutes, to about 20.7. Exposures of 5 or 6 minutes reached the extreme limits of the 100-inch, beyond the end of the sequence. From these data it is estimated that the 60-minute exposures permitted by the dark sky above Palomar reached at least 1.5 magnitude beyond the 100-inch. The threshold of the plate is dominated by nebulae rather than by stars, and this fact emphasizes the tremendous range of the telescope. Some of the faintest nebulae recorded are presumably at about twice the distance reached with the 100-inch or, in round numbers, at about 1,000 million light-years. This figure, of course, refers to average nebulae. Individual images on the photograph may represent dwarf nebulae at lesser distances or giant nebulae, even more remote. Plate 7, Messier 87 (NGC4486); R.A.=12°2873, Decl.=-+12°42’ (1950); Apr. 27, 1949; exposure 45 min., seeing average; enlarged 8X. The object is one of the brightest members of the Virgo cluster of nebulae, whose distance is of the order of 7.5 million light-years. It is classified as a peculiar elliptical nebula (KOp). The photograph shows the nebula, presumably a globular mass of type II stars (1. e., stars similar to those in globular star clusters), surrounded by an extensive, tenuous atmosphere of supergiant stars. ‘This phenomenon was suggested by the best photographs made with the 100-inch on Mount Wilson, but is conspicuous on the 200-inch plate. Plate 8, NGC5204; R.A.=13"28"0, Decl.=+ 58°38’ (1950); Jan. 31, 1949; exposure 30 min., seeing average; enlarged 4X. The object is a dwarf, late-type spiral in Ursa Major, at an esti- mated distance of less than 3 million light-years. The plate is in- cluded to show the ability of the telescope to resolve the neighboring stellar systems so that the brighter stars can be studied individually. Photographs of several of the larger spirals, such as M 81, NGC2403, etc., have been made, but the coma-free field of the telescope at full aperture is so small (about 5 minutes of are in diameter) that the plates are not suitable for reproduction on a scale sufficient to show the resolution to advantage. Plate 9, NGC3359; R.A.=10"43"4, Decl.=+63°20’ (1950); Apr. 27, 1949; exposure 45 min., seeing average; enlarged 3X. 188 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 This late-type barrel spiral is an isolated stellar system or, possibly, an outrider of the Ursa Major cloud of bright nebulae, and its distance is of the order of 5 million light-years. The plate is included to illus- trate the resolution of fairly distant nebulae, and the opportunities now available for the study of the very brightest stars in stellar systems. Plate 10, Messier 3 (NGC5272); R.A.=13539"5, Decl.=+28°38’ (1950); Apr. 21, 1949; exposure 3 min., made with an aperture of 160 inches, and a Ross correcting lens; seeing average; enlarged 8X. This plate, of a well-known globular star cluster, is included to illustrate the use of a Ross correcting lens, placed a few inches in front of the plate, to enlarge the coma-free field of the telescope. The lens performed well with the 160-inch aperture but, with the full 200-inch, it exaggerated the effects of the turned-up edge of the main mirror. The provisional mounting of the lens did not permit the use of a guiding eyepiece, so the plate shows the successful per- formance of the tracking mechanism of the telescope during an unguided exposure of 3 minutes. The usable field with this correcting lens is more than 15 minutes of are in diameter. THE DETERMINATION OF PRECISE TIME! By Str Haroup SPENCER JONES Astronomer Royal of Great Britain Of the three fundamental physical units, there is an essential distinction between the unit of time and the units of mass and length. The units of mass and length are represented by material standards to which any mass or length can be related, either directly or indirectly. But the unit of time cannot be represented by any material standard. For practical purposes time can be thought of in the Newtonian sense as something which flows uniformly. The passage of time can be marked by a clock, and any simple natural phenomenon which obeys one definite law without perturbation might be used to mark off equal intervals of time and therefore to serve as a clock. The rotation of the earth provides us with a natural clock. We shall see later that it is not a perfect clock, but that it is sufficiently uniform for almost all practical purposes; it has, moreover, the great advantage of never stopping. We can therefore define the unit of time as the period of rotation of the earth. Some reference object must be selected against which to measure the rotation. For the purposes of everyday life, time must be related to the sun, whose rising and setting gives the alterna- tion of daytime and nighttime. The day defined by the rotation of the earth with respect to the sun is called the true solar day; it is the interval between two consecutive transits of the sun across the meridian of any place. With this unit, true solar time is obtained by dividing the true solar day into 24 hours and calling the instant of meridian passage of the sun 12 hours. The time given by a sundial is true solar time. For practical purposes, however, true solar time is not convenient; because the motion of the sun across the heavens is not uniform, the length of the solar day varies in length throughout the year. For civil purposes, therefore, a mean solar day is used, whose length is equal to the average length of the true solar days throughout the year. The time based on the mean solar day as unit is called mean solar time. The relationship between mean solar time and true solar time at some particular instant is defined by means of 1 Sixteenth Arthur lecture, given under the auspices of the Smithsonian Institution April 14, 1949, 189 190 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 a convention, into the details of which I need not enter. The extreme differences between mean and true solar times range from 16 minutes about November 3, when true noon precedes mean moon, to 14% minutes about February 12, when true noon follows mean moon. Astronomers, however, find it more convenient to determine time by the observation of the stars. There are many stars but only one sun and, moreover, the time of transit of a star can be determined more accurately than the time of transit of the sun. The sidereal day is defined by the rotation of the earth relative to the stars. It is about 4 minutes shorter than the solar day. If we imagine the sun and a star to be on the meridian of some particular place at the same instant, then after the lapse of one sidereal day the star will again be on the meridian; but, because of the orbital motion of the earth round the sun, the earth will have to turn a little more in order to bring the sun onto the meridian. In the course of a year the earth completes its orbit around the sun and there must consequently be exactly one more sidereal day in the year than mean solar days. If the relative positions of a number of stars in the equatorial region of the sky have been accurately determined, we can think of them as equivalent to the graduations on the face of a clock. As the earth rotates, a telescope, fixed so as to be able to move only in the meridian, will sweep across these stars in turn, each at a definite specific instant of sidereal time. By observing the transit of stars whose positions are known, the sidereal times at the instants of meridian transit are therefore determined. The beginning of the sidereal day or, in other words, Oh. of sidereal time, is defined by the transit of the point in the sky at which the ecliptic crosses the equator from south to north; this point is called the vernal equinox or the First Point of Aries. By defining the commencement of the sidereal day in this manner, we are provided with a means for converting from sidereal time to mean solar time, which is required for the purposes of everyday life. But it has one inconvenience. The First Point of Aries is not fixed relative to the stars. It has a slow retrograde motion, due to the precessional motion of the earth’s axis, and superposed on this uniform motion is a slow to-and-fro drift, caused by the nutational or nodding motion of the axis. The nutation depends upon the relative positions and distances of the sun and moon from the earth. The principal term in the nutation has a period of about 18 years and a semi- amplitude of about 1 second of time. There is also a 6-monthly term amounting to 0.08 second and a number of short-period terms amounting to 0.020 second, of which the principal term has a period of 2 weeks. The precision of modern clocks is such that these small terms cannot be neglected. The true sidereal day, measured relative to the true position of the First Point of Aries, is therefore not abso- lutely uniform in length, and it is necessary to introduce the con- DETERMINATION OF PRECISE TIME—-SPENCER JONES 191 ception of mean sidereal time, measured relative to the mean position of the First Point of Aries. Actual observation of the stars provides the astronomer with true sidereal time, which he then has to correct for the nutation to obtain mean or uniform sidereal time. The determinations of time by astronomical observations are used to control the performance of a standard clock, determining its error at a specific instant and the rate of increase or decrease of that error, the clock then being used to obtain the time at other instants. This usually involves extrapolation to some time subsequent to the latest observation. For such extrapolation to be accurate, the time de- terminations must not be affected by serious errors and the standard clock must be of high precision. The determination of precise time therefore involves two problems, the determination with high ac- curacy of the time at specific instants and the development of time- keepers of very high precision. The sidereal time of the transit of a star across the meridian is equal to the right ascension of the star. Sidereal time can therefore be determined by observing the times of meridian transit of stars of known right ascension. The conventional method of making the observations has been to use a transit instrument. This consists of a telescope, mounted on an axis at each end of which is a cylindrical pivot. The pivots rest in fixed bearings, adjusted so that the common axis of the pivots is as nearly as possible horizontal and pointing in an east-west direction. If the axis of the pivots were exactly horizontal and in the east-west direction and if the optical and mechanical axes of the telescope coincided, the axis of the telescope would be in the meridian plane, whatever direction the telescope was pointing to. This ideal condition is never achieved and there are always small errors of level, of azimuth, and of collimation. These adjustments are liable to continual change; there are slow seasonal changes, as- sociated with changes of temperature and possibly also with sub- surface moisture; there are also more rapid changes, which are cor- related with changes of circumambient temperature and with the direction of the wind. To control these changes frequent observa- tions of level, of azimuth, and of collimation are essential, which take up a disproportionate amount of the observing time. The error of collimation can, however, be eliminated if the telescope is reversed in its bearings in the middle of each transit, half the transit being observed before reversal and the other half after reversal. It is not possible to reverse large transit instruments sufficiently quickly and it has accordingly become customary to use small transit instruments, which can be rapidly reversed, for the determination of time; as it is the brighter stars which are observed, a large aperture is not needed. There are other factors which have also to be taken into considera- tion. The pivots will never be absolutely cylindrical; their figures 192 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 have to be determined with great accuracy and appropriate corrections made to the observations. Flexure of the axis can cause troublesome systematic errors. If the horizontal axis is not equally stiff in all directions, its flexure will vary according to the direction in which the telescope is pointed. If the two halves are not equally stiff, the telescope will be twisted from the meridian by a variable amount. Personal equations between different observers are somewhat trouble- some, though they do not exceed a few hundredths of a second when the so-called impersonal micrometer is used. Before its introduction, the method of observing was for the observer to press a hand-tapper at the instant the star crossed each of a number of vertical spider wires in the focal plane of the telescope; by so doing, he closed an electric circuit which sent a current to a recording chronograph, which recorded not only the signals from the telescope but also time signals, every second or alternate seconds, from the clock. The instants of the star crossing the wires could then be read off at leisure after the observations had been completed. With this method of observing, the times determined by different observers could differ by as much as halfasecond. The reason is easy to see; one observer might wait until he saw the star actually bisected by the wire before he pressed the tapper, with the result that, because of the time required for the mes- sage to travel from his brain to his eye and to be converted into muscular action, his signal would inevitably be late; another observer would, as it were, shoot the flying bird, gauging the rate of motion of the star so that his tap is made as nearly as possible at the instant at which the star is actually bisected. The personal equations can be determined by what are called personal equation machines; the transit of an artificial star is observed, the times at which the star is at certain positions during the transit beng compared with the observed times. Although an observer will unconsciously form a fixed habit in observing so that his personal equation remains substantially constant, small variations, depending upon the physical condition of the observer, do occur. The method of observing now almost universally employed is to have a single movable wire in the micrometer eyepiece instead of a number of fixed wires. The wire can be traveled along by the observer, who adjusts its speed so as to keep the star continually bisected by the wire. As the wire moves along, contacts are auto- matically made in certain positions, sending signals which are re- corded on the chronograph. In order to relieve the observer of some of the strain of maintaining a uniform motion of the wire, it is now common to drive the wire mechanically at the speed appropriate to the motion of the star, using an electric motor with some form of con- tinuously variable gearing. With this method of observing, the personal equations of different observers are very small, usually not DETERMINATION OF PRECISE TIME—SPENCER JONES 193 more than two or three hundredths of a second; it is for this reason that this form of micrometer is called the “impersonal”? micrometer. Small though these residual personal equations are, they remain remarkably constant and can be determined by personal equation machines. They seem to arise from two causes: there is “bisection error,” an observer systematically bisecting an image to the right or to the left of its center; this error changes sign at the zenith with instruments in which the observer changes the direction in which he faces, according to whether he is observing a north or a south star; there is also “‘following error,” an observer systematically setting the wire in front of or behind the center of a moving image. This error does not change sign at the zenith. If the pivots are not exactly cylindrical, the telescope will be twisted out of the meridian by an amount varying with its position. The figures of the pivots must therefore be determined with great accuracy and appropriate corrections applied to the observed times of transit. The figures of the pivots must be determined at intervals, as they may change slowly in the course of use through wear. Other variable errors can be introduced through slight mechanical imperfec- tions in the telescope; if there is the slightest play in the eyepiece micrometer or in the objective, errors will be introduced which will vary with the position of the telescope. When all the possible sources of error which can affect observations with a transit instrument are borne in mind, it is rather surprising that the observations are as accurate as they are. The probable error of a single time determination is usually about two-hundredths of a second. This was quite accurate enough before the era of clocks of high precision and before there were any practical requirements for very precise time. The scatter of the observations is, however, incon- veniently large for the adequate control of the performance of the modern quartz-crystal clock. For these reasons the conventional transit instrument is likely to be gradually replaced for the purpose of time determination by some other type of instrument. Several modifications of the transit instrument have been considered which eliminate or minimize some of its disadvantages. The most accurate results are given, however, by an entirely different instrument known as the photographic zenith tube. It consists of a fixed vertical telescope pointing to the zenith, which has a mercury horizon at the bottom of its tube, whose purpose is to reflect the light from a star to a focus in the plane of the second principal point of the objective. The funda- mental principle of the instrument was due to Sir George Airy, who first used it for the reflex zenith tube at Greenwich: when the light is brought to a focus accurately in the plane of the second principal point of the objective, the results are unaffected by tilt of the telescope. The troublesome error of level is therefore immaterial, while any error 194. ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 of azimuth does not affect observations made in the zenith. The telescope is constructed so that the objective and the plate holder can be rotated through 180°, the observations being made photographically in order to eliminate personal equations and to give greater accuracy. Suppose two exposures are given on a star at times which are sym- metrical about the time of meridian transit, the objective and the photographic plate being rotated through 180° between them. The two images will lie on a line exactly parallel to the meridian. If, however, the two times of exposure are not exactly symmetrical, the images will be slightly staggered; by measuring the staggering and knowing the clock times of the two exposures, the clock time of meridian transit can be inferred. In practice an exposure of finite length is required to give a measurable image on the plate. During this exposure the plate holder is traveled along at the speed appropriate to the motion of the star, signals being sent to the chronograph at certain definite positions of the plate holder. After reversal the plate carriage retraces its path, and signals are sent during the course of the second exposure at the same positions. With this design of instrument, collimation error does not enter, there are no pivot errors to be considered, and the various sources of error inherent in a movable instrument are avoided. At the Naval Observatory, Washington, a photographic zenith tube, designed and used by F. E. Ross originally for the determination of the variations of latitude, has been used for some years for the determination of time. An instrument on the same general principle, but differing materially in details of design, is in an advanced stage of construction for the Royal Greenwich Observatory. The errors of time determination should not exceed 2 or 3 milliseconds, which will permit a tight control of the performance of the observatory clocks. For the purpose of time determination it is necessary to assume positions for the stars which are observed. These positions will have random errors, whose effects can be reduced by observing sufficient stars. But they may also be affected by systematic errors; if, for instance, the errors vary with right ascension they will introduce a spurious systematic variation in the derived clock error through the year. For the purpose of time determinations and in order that the times determined at different observatories can be directly compared, there is an international agreement to use the positions of the stars given in the fundamental star catalog known as the FK3. These are bright stars, whereas with the photographic zenith tube, inasmuch as observations are restricted to a narrow belt at the zenith, it is necessary to use fainter stars. Their positions must therefore be determined by transit circle observations and tied on to the FK3 system, The photographic observations will in course of time provide DETERMINATION OF PRECISE TIME—SPENCER JONES 195 some measure of control over the periodic errors in right ascension of the FK3 system itself. Until about 25 years ago, pendulum clocks of the regulator type were used as the standard clocks in observatories. A considerable improvement in precision was brought about by the invention of the free-pendulum clock. In an ordinary pendulum clock the timekeeping is impaired by the variable friction involved in driving a train of wheels to move the hands and record the actual time on a dial. An appreciably higher accuracy is to be expected if the pendulum is allowed to swing freely, except when it receives periodically impulses to maintain its swing, and is thereby relieved from all extraneous work. To achieve this had been the aim of horologists for many years, but although many attempts were made it was not really successfully accomplished until the invention by W. H. Shortt of his free-pendulum clock. The master pendulum is enclosed in an airtight case, in which the air pressure is reduced to about 1 inch of mercury and which is maintained at constant temperature and swings freely, except for small impulses, given at half-minute intervals, to main- tain the amplitude at a nearly constant value. The slave clock is a normal synchronome electric clock, which is adjusted when swing- ing as an independent clock to lose about 6 seconds a day. The synchronizing action required from the master free pendulum is there- fore a one-way action—always an accelerating action. The slave pendulum itself releases electrically the impulsing lever of the free pendulum, which falls when the free pendulum is at the midpoint of its swing. The impulse arm falls on the top of a small pivoted wheel, mounted on the free pendulum; this being a dead point, and the impulse not commencing to be given until the pendulum swings outward from the central position, the amount of the impulse does not depend upon any slight variation in the synchronization between the two pendulums which may occur. The synchronization of the slave pendulum is effected by means of a light flexible spring carried on it. The impulse arm of the free pendulum, after it has fallen clear of the pendulum, actuates a device which closes an electric circuit and sends a current through a small electromagnet adjacent to the slave pendulum. If the slave clock has dropped sufficiently behind the master, the armature of this electro- magnet will, when the electromagnet is excited, engage the bent end of the light spring on the slave pendulum. The end of the spring is then held fixed and, as the pendulum swings, the spring is flexed and the pendulum is accelerated; if, on the other hand, the slave pendulum is closely in phase with the master, the end of the spring passes under the armature before the electromagnet is excited, and nothing happens. The length and strength of the spring are so adjusted that when the synchronizing action occurs the slave pendulum is accelerated by 196 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 %4o second. As the natural losing rate of the slave clock, 6 seconds a day, is equivalent to go second per minute, the synchronizer, which is actuated each half minute, should hit and miss alternately. For this reason it is called the ‘‘hit-and-miss” synchronizer. The first experimental Shortt free-pendulum clock was installed at the Edinburgh Observatory in 1921. It at once proved to be such an improvement upon previous pendulum clocks that two were installed at the Greenwich Observatory in 1923 and others in subsequent years. It was the excellent performance given by these clocks that made it necessary for astronomers for the first time to introduce the conception of mean sidereal time. Previously true sidereal time had been uni- versally used, as clocks were not good enough to be able to show up the small effects due to the short-period terms in nutation. The free- pendulum type of clock is capable of an accuracy of about one- hundredth of a second a day. Detailed investigation of their performance has shown, however, that such clocks are liable to frequent small erratic changes of rate, of the order of about 3 milli- seconds a day. Small though such changes are, they cause, by integration, an irregular wandering of the clock. For sending out time signals, it is always necessary to extrapolate beyond the latest time determination; these erratic changes of rate restrict the accuracy with which the error of the clock can be extrapolated. It can, on occasion, happen that 2 weeks or more may elapse without any check on the performance of the clock being possible and the transmitted time signals may consequently be appreciably in error. Moreover, because of the errors of observation, there is a natural scatter in the derived errors of the clock. In interpolating between the observed errors there is no means of distinguishing between scatter due to errors of observation and scatter due to the irregular wandering of the clock. It is possible, of course, to attempt to reduce the effects of the wandering by using the mean of several clocks. Nevertheless, very high accuracy cannot be obtained, because residual effects due to the irregularities are always present. A new standard of accuracy has been provided in recent years by the use of an oscillating quartz crystal, developed originally to serve as a precision standard of frequency. The quartz clock is based upon the piezoelectric property of quartz. If a plate of quartz is com- pressed, the two opposite faces become electrically charged, one positively and the other negatively. Conversely, if two opposite faces are given positive and negative charges respectively, the piece of quartz experiences a mechanical contraction or expansion. By rapidly alternating the electric charges, the quartz can be maintained in mechanical vibration. In the quartz clock an oscillating electrical circuit is used, the dimensions of the crystal being adjusted so that its DETERMINATION OF PRECISE TIME—SPENCER JONES 197 natural resonance frequency is equal to the frequency of the oscillating circuit. Under these conditions a strong vibration is set up and the quartz crystal takes control and locks the frequency of the oscillating electrical circuit to its own resonance frequency. Quartz is a very stable substance and, provided it is maintained at a very uniform temperature and the drive circuit is properly designed, the frequency remains constant to a high degree of accuracy. It is usual for the crystal to be cut to give a frequency of 100,000 cycles a mean time second, the dimensions of the quartz then being conveniently small. This frequency is divided down in steps electronically, either by the use of multivibrators or by frequency subdivision until an output with a frequency of 1,000 cycles a second is obtained. The output of this frequency is used to drive a phonic motor, from which time signals can be obtained at any desired intervals. Such clocks have many advantages over pendulum clocks. They have proved to have very high short-period stability. Their erratic changes of rate are less than half a millisecond a day, and the clocks themselves can be relied upon to about 1 millisecond a day. For extrapolating between scattered time determinations they are there- fore much superior to pendulum clocks. They have, moreover, the advantage of the great flexibility inherent in dealing with 100,000 vibrations a second instead of only a single one. Electronic methods can be used for quickly and accurately determining the relative errors and rates of the clocks. For such purposes at Greenwich, decimal counter chronometers are used. This device consists of a scale-of-ten counter, and is actuated by the 100,000-cycle output per second from one of the quartz crystals. When it is switched on, it will start counting these vibrations, recording the count on five decade dials, reading, respectively, tenths, hundredths, thousandths, ten- thousandths, and hundred-thousandths of a second. To compare two quartz clocks, a seconds signal from the phonic motor driven by the one clock is used to start the count and a signal from the second clock to stop it. The time difference between the two clocks, accurate to a hundred-thousandth of a second, is thus obtained in a fraction of a second. As a check, the second clock can be used to start the count and the first clock to stop it. The difference in frequency of the two clocks is obtained by feeding the 100,000 ¢. p. s. outputs from the two clocks into a comparator, so that they beat against one another, and timing the beats. It is possible to obtain an accuracy of one part in 10”° in the measurement of the frequency difference. At Greenwich, the clocks are used in groups of three, one phonic motor being provided for each group of three clocks. One of the clocks is selected to drive the phonic motor, but regular comparisons are made between each pair of clocks in the group. Automatic beat 198 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 counters record the integrated time difference between each pair, A-B, B-C, C-A, the third comparison providing a check on the other two. A further convenience of the quartz clocks is that it is not necessary to maintain separate mean-time and sidereal-time clocks as it is with pendulum clocks. By means of suitable gearing, it is possible to take sidereal seconds direct from the phonic motor which gives also mean time seconds. The ratio of the mean time second to the sidereal time second is 1.002 737 909 293. This ratio can be closely represented by a gearing of 119/114 multipled by 317/330, which is only 4 parts in 10° small. These sidereal second signals are used for recording on the chronograph during the time determinations. When the rate of the clock relative to these signals has been derived it is a simple matter to infer its rate relative to true mean time seconds. The small error in the conversion from mean time to sidereal time is, of course, eliminated. For short-period prediction quartz clocks leave little to be desired. They have not as yet, however, reached the stage at which long-period prediction has the accuracy that is desirable. The difficulty arises from a slow drift in frequency to which they are all liable. The crystal, after cutting, appears to go through a slow ageing process; the drift in frequency is rather rapid at first, but progressively diminishes though it sems never to cease altogether. Iffor any reason the crystal should stop, through a tube or resistor giving out, it will not, when restarted, follow along its previous ageing curve; a new ageing cycle sets in. Any small disturbance, such as a slight temperature change, can alter the frequency drift somewhat. The effect of the frequency drift on the error of the clock increases with the square of the time so that, even though the drift may be quite small, its effects will become important with lapse of time. With the present scatter in the actual time determinations, several months’ observations are needed to give a sufficiently accurate derivation of the frequency drift, but there is always the uncertainty whether during this period some small disturbance may not have caused the rate of drift to change slightly. Moreover there are extraneous effects which can complicate the determination. During a period of several months, there will be a wide range in the right ascensions of the stars which are used for the time determinations. If there are periodic errors in the fundamental system of star places, a spurious factor will have entered into the determination of the frequency drift. The motions of the earth’s poles cause further complications. The poles have an irregular mo- tion, which is roughly circular, but with a variable radius. The extreme departures of the true poles from their mean positions are about 30 feet. ‘The movement of the pole along the meridian causes DETERMINATION OF PRECISE TIME—SPENCER JONES 199 a variation in latitude, which can be observed with a zenith tele- scope. The movement in the perpendicular direction causes a dis- placement of the meridian. The motion of the pole has two main components, with periods of a year and of about 14 months respec- tively. As a consequence of this motion, it would be found that if we had a perfect clock, with no rate at all, and observations which were entirely free from error, the clock would appear to have a slightly variable rate. This apparent variation of rate will affect the deter- mination of the frequency drift and give a spurious value. It is not possible at an observatory to measure the component of the polar motion at right angles to the meridian. At Greenwich an approximate compensation for the motion is made through the cooperation of the Naval Observatory, Washington, which sends regularly to Greenwich the observed movement of the pole along the meridian of Washington. If Washington were 90° in longitude west of Greenwich, the displacement along the meridian of Washington would also be the displacement at right angles to the meridian of Greenwich. But the longitude of Washington is only 77° west of Greenwich. However, the use of the Washington latitude-variation data does enable the greater part of the polar-motion effect to be elim- inated from the Greenwich clock curves and it has been noticeable that the inferred performance of the clocks has thereby been improved. The development of an atomic or molecular clock, in which the frequency of some selected atomic or molecular vibration will be subdivided to give a frequency closely equal to that of an oscillating quartz crystal and used to lock the vibrations of the crystal, is already foreshadowed by the work in progress at the National Bureau of Standards, Washington, in the development of an ammonia clock, in which the frequency of one particular mode of vibration of the ammonia molecule is used as the control. This work is as yet in its early stages and has not gone beyond the point of showing that the control of a quartz crystal in the way suggested is practicable. When the clock has been developed to the stage at which the accurate con- trol of a precision quartz clock becomes possible, the crystal will be prevented from drifting in frequency. ‘The clock error curve over a long period of time should then be represented by a straight line. Departures from a straight line could be attributed to periodic errors in the star places, to the polar motion, or to irregularities in the rate of rotation of the earth itself. Much more accurate long-term prediction would become possible, with a considerable gain in the accuracy of timekeeping. It has been well established that the length of the day is subject to small fluctuations. It has long been known that there are discord- ances between the observed and the tabular positions of the moon which are not attributable to imperfections in the theory of the 866591—50-——14 200 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 motion of the moon. In the development of the theory, the gravita- tional effects which have been neglected are far too small to amount to anything like the discordances which are observed. In more recent years it has been proved that there are similar fluctuations in the motions of Mercury, Venus, and the sun; but for these bodies the effects are much smaller than for the moon because their mean motions are much less rapid. It was the comparative smallness of the effects for these bodies which made their detection difficult. So there are, in effect, four clocks which agree together and one clock, our earth, which differs from the other four. The natural conclusion is that it is the earth which is at fault and that the length of the day, which has been adopted as the unit of time and assumed to be invar- iable, is actually subject to small variations. The changes in the length of the day are found, from the analysis of the observational data, to be of two different kinds. There is a slow progressive increase in length, of the order of 1 millisecond in the length of the day in the course of a century. This progressive increase is caused by tidal friction, more particularly in the shallow sea; it acts as a brake on the earth. Though so small in amount, the effect on the mean longitudes of the moon and the planets increases with the square of the time and is large enough to make the position of the moon 20 centuries ago, if computed from its present motion in longitude, very considerably in error. The effect was actually first detected in 1679 by Hailey from the early observations of eclipses. Superposed on the progressive increase of length there are also irregular changes, the day sometimes increasing in length and sometimes de- creasing; these changes cannot be attributed to tidal friction, because frictional effects can cause only a slowing down and never a speeding up in the earth’s rotation. These changes are due to changes in the earth’s moment of inertia and could be accounted for quantitatively if the earth expanded or contracted slightly by 4 or 5 inches. There is one essential difference between the two phenomena. A change in the moment of inertia of the earth is something that con- cerns the earth alone. The apparent displacements of all the other bodies are strictly proportional to their mean motions. But tidal friction is something that concerns the earth and the moon jointly; the total angular momentum of the earth-moon system is conserved, but there is interaction between the earth and the moon. The appar- ent displacements of Mercury, Venus, and the sun will again be proportional to their mean motions but the same will not hold for the moon; its displacement will not have the same ratio to its mean motion. It is this difference in the case of the moon which makes it possible to separate the two effects of tidal friction and of change of the moment of inertia of the earth. DETERMINATION OF PRECISE TIME—SPENCER JONES 2(1 Though the changes in the length of the day have been fully estab- lished by these observations, the data are not sufficiently accurate to decide whether the changes occur suddenly or whether they are spread over a few days, a few weeks, a few months, or even over a year or two. If they occur rather suddenly, they could be detected with ease by quartz clocks in their present stage of development; if spread over a few months, the larger changes could be detected, but changes of smaller amount would be likely to escape detection. Since a few years ago, when quartz clocks were adopted at Greenwich as the basis for the time service, a close watch has been kept for any evidence of a change in the earth’s rotation. Once or twice small changes have been suspected but there has always been some factor which has made a definite conclusion impossible—perhaps one of the clocks has changed its rate or has stopped at the crucial time, or there has been some uncertainty in the determination of frequency drift. The evidence provided by the observations of occultations of stars by the moon is that there has been no major change in the earth’s rate of rotation since about 1918. There may possibly have been small changes, but no definite conclusions are as yet possible. It is not inconceivable that there may be small annual variations in the rate of rotation of the earth. There are seasonal displacements of matter over the earth’s surface; there is, for instance, a high-pressure region over Siberia at one season of the year and a low-pressure region at another season, entailing the displacement of large atmospheric masses, with corresponding change in the moment of inertia. Such effects would be tangled up with effects due to periodic errors in star places and with the effects of the polar motion. Much more is likely to be learned about these matters when the atomic clock has reached a further stage of development, so that the frequency drift of the quartz crystal can be eliminated. Observations with photographic zenith telescopes should gradually smooth out any residual periodic errors in star places, while the information they provide about the variation of latitude will furnish basic data which can be used subse- quently to separate polar motion effects from small variations in the earth’s rotation. It may prove, however, that the earth itself is rather like a pendulum clock in its behavior and that its rate of rota- tion is liable to frequent and small irregular changes, so that we can at present merely observe their integrated effect. The question may arise in the near future how the unit of time should be defined. Clocks are now at a stage when their stability for short periods is of a higher accuracy than the earth’s rotation itself. The earth, however, has the advantage over any clock that it has no liability to a stoppage. It may be possible to develop atomic clocks to a stage at which they can be run for several years 202 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 without stopping and to maintain accurate time; with a battery of such clocks, all controlled by the same atomic vibration, it would be possible to bridge over the stoppage of any single clock and thereby to maintain an accurate standard of time more or less indefinitely. There will be definite objections to using as the fundamental unit of time a unit that is known to be variable. A new unit should be absolutely invariable. A clock based fundamentally upon a length which is controlled by an atomic wave length, and upon the velocity of light, for instance, seems theoretically ideal. Note ‘ADDED IN PROOF.—Since the lecture was delivered, investi- gations at the Greenwich Observatory have established the existence of a fairly regular annual variation in the rate of rotation of the earth. Relative to uniform time the earth gets behind by about 60 milliseconds in May-June and ahead by a similar amount in Novem- ber. The corresponding variations in the length of the day amount to somewhat more than 1 millisecond a day on either side of the mean value. H.S. J. THE ELEMENTARY PARTICLES OF PHYSICS! By Cart D. ANDERSON California Institute of Technology, Pasadena The idea of elementary particles of matter, of small, discrete, in- divisible particles out of which all matter in the universe is consti- tuted, is as old as recorded history. The Greeks in their philosophical speculations discussed at length the question of the ultimate nature of matter. They realized that there were only two possible choices open to them; either matter must be thought capable of being divided into smaller and smaller units without end, or else it must consist of small units which are themselves wholly indivisible. Many of the Greek philosophers experienced a philosophical difficulty in trying to conceive of infinite divisibility, whereas others found it equally difficult to think of a particle as being truly indivisible. The difficulty is closely akin to that which one experiences when contemplating the limits of the uni- verse, and trying to decide in his own mind whether it pleases him more to think of the universe as unbounded and extending to infinity, or to imagine a finite universe with definite bounds beyond which there is nothing, not even space. The idea of the existence of indivisible ma- terial particles, however, seems to have had more appeal to the Greeks, and the atomic hypothesis was expounded and developed in the fifth century B. C., chiefly by Thales, Leucippus, and his distinguished pupil, Democritus, until in many respects it resembled the views which are held today. The views of Democritus were prominent for 500 years but began to wane after the beginning of the Christian Era and by about A.D. 200 had almost wholly disappeared from European philosophical thought. The idea of material atoms did not really appear again in Europe until about the middle of the seventeenth century, a time marking the beginning of the great era of scientific experimentation which has continued with an ever increasing tempo up to the present. During this period, through scientific research based on experimen- tation, the atomic theory of matter slowly developed. Highlights in 1 Based on material presented in the Sigma Xi Annual Address, A. A. A. S. Centennial, Washington, September 1948. Reprinted from American Scientist, vol. 17, No. 2, April 1949, by permission from The Society of the Sigma Xi. 203 204 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 this development were the Laws of Chemical Proportion as discovered and enunciated by Dalton near the beginning of the nineteenth cen- tury, and later the successes of the Kinetic Theory of Gases. By the beginning of the twentieth century, the concept of the chemical atom had received general acceptance as a theory based on scientific experi- mentation. The idea of atoms had thus been removed from the realm of philosophical speculation and had become a proved scientific fact. According to this picture all matter depending upon its nature consists of a mixture of varying numbers of the ninety-odd different chemical atoms. The size and the mass and other properties of most of the chemical atoms had been determined although not with great preci- sion. DISCOVERY OF FIRST ELEMENTARY PARTICLES During the time when the chemical atom was being firmly estab- lished as a scientific fact, other scientific investigations were succeed- ing in proving the existence of at least one particle of matter which was more elementary in character than the chemical atoms. In the decade from 1890 to 1900 the discovery of X-rays and radioactivity, and studies of the phenomena associated with the discharge of elec- tricity through gases, soon proved the existence of the electron and showed that the atoms of chemistry must all be considered as complex structures, structures which are themselves built up of particles of a more elementary character. The electron was distinguished from the other particles previously studied by physicists and chemists in one very important respect. It was established as a unique particle in the sense that all electrons were found to be identical with one another, no matter from what form of matter they were derived. For the first time then the presence of a particle truly elementary in character was revealed to science. It was found always to carry a negative electric charge and to have a mass about 2,000 times less than the hydrogen atom, the simplest and least massive of all the chemical atoms. The electron immedi- ately took its place as one of the elementary particles common to all forms of matter. The following 30 years, from 1900 to 1930, were extremely fruitful in furthering our knowledge of the properties of the chemical atoms. The work of Moseley showed that chemical atoms were members of a family, all of them being related to one another in a perfectly definite and simple way. In 1911 the experimental genius of Rutherford in Cambridge, England, proved the existence of the atomic nucleus, and in 1919 he succeeded for the first time in producing an atom of oxygen from the disruption of the nucleus of an atom of nitrogen. Thus in 1919 the will of man for the first time was able to cause the disinte- gration of an ordinarily stable element, with the accompanying release ELEMENTARY PARTICLES OF PHYSICS—ANDERSON 205 of nuclear energy. These and other investigations all combined to prove that the proton, the nucleus of the simplest of all the chemical atoms, hydrogen, is a constituent of all other chemical atoms, and hence is in fact one of the elementary particles of matter. In 1930, then, the physicist had at his disposal two elementary ma- terial particles, the electron and the protron, in terms of which to try to understand the structure of all matter. In this undertaking the physicist realized many great successes, but in many instances his efforts resulted in sharp failures. Apparently the world was not to be understood in terms as simple as these. In general the physicist was successful in understanding those phe- nomena which we may classify, for want of a better term, as extra- nuclear phenomena, and he was unsuccessful in understanding those phenomena which we may classify as nuclear phenomena. By extra- nuclear phenomena we mean those processes in which the electrons which form the outer shells of the atom are the active participating agents; in this type of phenomena the central core of the atom, or the nucleus, is present but remains undisturbed and does not participate actively. Nuclear phenomena, on the other hand, are those in which the nucleus is the active participant. Extranuclear phenomena and nuclear phenomena have a great many distinguishing characteristics. One of the most interesting and important of these distinguishing characteristics is concerned with the level of energies involved. Extranuclear phenomena involve very low energies as compared with nuclear phenomena. The physicist uses the term electron volt as a measure of energy. The energies of extranuclear phenomena are found usually to range from a fraction of one electron volt to several electron volts, whereas nuclear phenomena are found usually to correspond to several millions of electron volts. In our environment almost every phenomenon in nature represents an extranuclear phenomenon: for example, the burning of coal, the growth of plants, the generation of electric power by conventional means, the fermentation of wine, the explosion of dynamite, and others in uncountable numbers. Nuclear phenomena are not so common- place, but a few examples may be mentioned: for example, the gen- eration of the sun’s heat, the decay of radium, the manufacture of plu- tonium, the absorption of cosmic rays in the earth’s atmosphere, the explosion of an atom bomb. The concept of energy has been introduced here because of the great importance that this concept has in the discussion of any physical phe- nomenon. I have stated that extranuclear phenomena represent low- energy phenomena and nuclear phenomena represent high-energy phenomena. To be more accurate I should have said that in extra- nuclear phenomena we find low concentrations of energy; that is, the energy changes that one associates with a single elementary particle 206 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 are low in extranuclear phenomena and high in the case of nuclear phenomena. Moreover, physicists for the past several years have been studying certain phenomena which represent energy concentrations many thousands of times greater than those represented even by nuclear phenomena. This range of energies has been called the range of ludicrously high energies. So far the only opportunity the physicist has had to study phenomena in the range of ludicrously high energies is in connection with observations associated with cosmic rays, and we shall see in a moment that important knowledge of the elementary particles of matter has come from studies of phenomena in the range of ludicrously high energies. As stated previously, by 1930 two elementary particles of matter were known to the physicist, the electron which always occurred with a negative electric charge and the proton which always occurred with a positive charge. When considered in a manner consistent with the theoretical concepts as they had been developed up to that time in terms of the quantum mechanics, the negative electron and the positive proton served quite successfully as building blocks in terms of which to understand the structure of atoms so far as the extranuclear phe- nomena were concerned. But when attempts were made to picture the structure of the nuclei of the various chemical atoms, or to understand nuclear phenomena, the attempts usually ended in failure. Then suddenly in 1932 two new elementary particles were dis- covered: the neutron and the positive electron, or positron. The known elementary particles were therefore doubled in number, increasing from two to four, and providing the physicist with more material with which to work. The discovery of the neutron, which came as a result of experiments performed in Germany, in France, and in England, was immediately welcomed, for now neutrons together with protons could serve as the building stones for the various types of atomic nuclei. One very grave and fundamental problem which formerly had been present was now removed immediately, for it was no longer necessary to assume the existence of electrons inside the nucleus, a concept which always had been accompanied by serious theoretical difficulties. The discovery of the positive electron, or positron, came during a series of experiments being performed for the purpose of measuring the energies of the particles produced by cosmic rays. The discovery of the positron was an unexpected discovery. This statement is true even though, about 2 years before, a British physicist, Dirac, had an- nounced a new theory which actually predicted the existence of posi- trons. This new feature of physical theory was not welcomed by physicists, however; it was on the contrary considered to be an un- fortunate defect in the theory, and many attempts, by Dirac himself ELEMENTARY PARTICLES OF PHYSICS—ANDERSON 207 and others, were made to remove it, although all were unsuccessful. If even one physicist in the world had taken the theory of Dirac seriously, he would have had an admirable guide leading directly to the discovery of the positron. Had this happened, the positron would almost certainly have been discovered by 1930 rather than in 1932. However, after the positron was shown actually to exist, then it was a very short time indeed until many of its properties were understood in terms of the Dirac theory. ELEMENTARY PARTICLES AND RADIATION The discovery of the positron represented the first instance in which it was recognized that an elementary particle of matter may have only a transitory existence. In ordinary matter, for example, the average life span of a positron is only a few billionths of a second, for when a positron and a negative electron come close to one another they mutually annihilate one another—the two particles disappear and in their place one finds only radiation. The whole of the material sub- stance constituting the particles is spontaneously transformed into radiant energy. Measurements show that this process is quantita- tively in accord with the now famous Einstein equation H=mc?, which relates mass and energy. ‘The process which is the inverse of the annihilation of material particles also occurs, namely, the production of particles out of radiation. If radiation of sufficiently high energy is passed through matter, electrons and positrons are generated. In this process the material substance of the two particles is actually created out of the energy represented by the radiation, and again in conformity with the Einstein equation H=me’. In the light of these happenings one must change basically his con- cept of the elementary particles of matter; these particles are no longer to be thought of as permanent objects which always preserve their identity, and which serve only as building blocks of matter by joining together in groups to form the more complex chemical atoms. One must recognize instead the possibility of the creation of material particles out of radiation, and the annihilation of material particles through the production of radiation. Such a possibility as this, of course, was completely inconceivable to the Greeks in their long philosophical discussion on the indivisibility of matter versus the divisibility of matter. A further step toward a realization of the great complexity inherent in the relationships among the elementary particles of matter came in 1935 with the discovery of the positive and negative mesotrons, or positive and negative mesons as they are now often called. This dis- covery was also made in investigations of the high-energy phenomena occurring when cosmic rays are absorbed in their passage through matter. 208 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The mesotron is a particle some 200 times as massive as an electron, and therefore about one-tenth as massive as either a proton or neutron. It occurs with both positive and negative electric charge. The dis- covery of the mesotron did not come quickly and accidentally as was the case with the positron and the neutron. It came only after the completion of a sustained series of observations, covering a period of 4 years, which were designed to remove certain inconsistencies always present when we attempted to understand certain ‘cosmic-ray phe- nomena in terms of the elementary particles then known. These inconsistencies were removed in terms of the existence of the mesotron, whose discovery was publicly announced in 1936. Unlike the neutron, the mesotron was not a particle to be imme- diately welcomed by the physicist. The physicist makes his advances by simplifying his understanding of nature; hence, a physical world which could be explained in terms of only one or two distinct elemen- tary particles would be most to his liking. The discovery of the mesotron did not introduce a simplification; rather it complicated the situation for it increased the number of material elementary particles from four to six. Apparently the Creator does not favor a world of too great simplicity. Before the discovery of the mesotron a Japanese physicist, Yukawa, had postulated on theoretical grounds the possible existence of parti- cles of a mass intermediate between a proton and an electron. His theory, however, was not generally known to physicists at that time, and did not have any part at all in the discovery of the mesotron. Had this theory been generally known it is still doubtful if it would have affected the course of cosmic-ray research, since, unlike the Dirac theory of the positron, it would not have served as so useful a guide in pointing out the most fruitful directions for the research to follow. Like the positron the mesotron has a very short life expectancy. In free space, both positive and negative mesotrons have a normal life span of just over two-millionths of a second, after which time they spontaneously disintegrate. Very recent observations have shown that in all probability the spontaneous disintegration of a mesotron results in the simultaneous production of an electron and two neutrinos. Neutrinos are the interesting elementary particles which had pre- viously been invented in order to balance energy and momentum in the process in which an electron is produced when a radioactive nucleus decays. A similar situation exists in the case of the decay of a mesotron except that here, because the mesotron disappears entirely, it is necessary to postulate the emission of two neutrinos in order to balance energy and momentum. In free space mesotrons spontaneously decay after about two- millionths of a second. In the presence of matter, a mesotron of ELEMENTARY PARTICLES OF PHYSICS—ANDERSON 209 negative charge may terminate its existence in an even shorter time. It does this by entering an atomic nucleus or, in the language of the physicist, by undergoing nuclear capture. The mesotrons observed in cosmic rays are produced by the very high energy particles of the primary cosmic-ray beam as it comes into the earth from outer space and plunges through the earth’s atmos- phere. In a manner somewhat analogous to the creation of positrons and electrons, the mesotrons are born out of the tremendous energies carried by the primary cosmic-ray beam. There are many interesting phenomena involved in the birth and death of mesotrons and in the violent nuclear processes which accom- pany these phenomena, but it will not be possible to discuss them here. However, I should like to mention in this connection two important advances which have been made within the last 2 years. RECENT ADVANCES IN NUCLEAR RESEARCH One of these is the work under way in Bristol, England, by Powell and his coworkers, which has consisted of a detailed analysis of the tracks produced by mesotrons in the emulsions of photographic plates. These investigators have discovered a mesotron of a new type which is heavier than the ordinary mesotron. It is about 285 times as massive as an electron, whereas the ordinary mesotron is about 215 times as heavy. The heavy mesotron has only a very short life; it lives only about one one-hundredth as long as the light mesotron, after which time it disintegrates and produces a light-weight mesotron and another particle which is probably a neutrino. The negatively charged heavy- weight mesotron may also directly enter an atomic nucleus and give rise to a violent nuclear disruption. Although both the newly discovered heavy mesotrons and the light mesotrons discovered in 1936 have some properties in common—e. g., both types of particles occur with positive and negative charges, both have short lives, and both are found in cosmic rays—nevertheless in some very fundamental respects they are entirely different types of elementary particles. The heavy mesotron interacts very strongly with atomic nuclei, but the light mesotron interacts only very weakly with atomic nuclei. Another difference lies in the respective values of that important property known as the spin or angular momentum; recent researches indicate that the heavy mesotron has an integral spin, whereas the light mesotron has a half-integral spin. In all probability it is the heavy mesotron and not the light mesotron which is to be identified with the particle first postulated on theoretical grounds by Yukawa in 1934. The theory of Yukawa even in its pres- ent state today is very primitive. However, this theory still provides the best basic concept in terms of which to understand processes in- volving mesotrons, and after further development in the future the 210 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Yukawa theory may possibly provide an understanding in terms of mesotron exchange forces of that all-important problem as to the nature of the forces acting between the particles inside a nucleus. So far no satisfactory theory has been developed in terms of which to understand many of even the simplest phenomena involving the nucleus. To acquire a quantitative understanding of the interactions of the elementary particles of matter and of the fundamental nuclear processes is one of the great tasks of theoretical physics today. To complete our list of elementary particles we should also include the photon. This particle, together with the neutrino as noted above, is, however, in a somewhat different category from the other types of particles. The photon is not a material particle, in the sense that it cannot be identified with any particle which can exist at rest and have associated with it a finite amount of ponderable material sub- stance. Photons are to be identified only with radiation or radiant energy. ‘The neutrino must also be placed in a special category, since it cannot have associated with it an appreciable amount of ponderable material substance if any at all, and since it has never been directly observed. Tn all, then, the physicist at the present time recognizes at least 10 distinct elementary particles of matter. Whether this list is complete or not no one can say with certainty. The indications are that the list is not complete, for evidence seems to be rapidly accumulating for the existence of at least one additional elementary particle. This particle is found in cosmic rays and appears to have a mass some 1,000 times the mass of the electron. But what its properties are and how it is related to the light and heavy mesotrons and to the other elemen- tary particles of matter is a subject which must await the results of further observations. The thought of probable further additions to the list of elementary particles of matter suggests a question which is quite apart from physics and has to do simply with the naming of new particles. We have here actually an interesting example of the great difficulties that physicists sometimes have merely in assigning labels or names to the various concepts which their experience or their theories have brought forth. It is usually necessary to choose some sort of name for these concepts, whether they be elementary particles of matter or something else, at a time before all the facts regarding them are known. In 1937 the term mesotron was suggested to designate the new particle of intermediate mass discovered in the cosmic rays in 1936. Since then this term has often been contracted to meson and has been so em- ployed. Since the discovery of the new particle whose mass is greater than the mass of the original cosmic-ray mesotron, the term mesotron or meson has been employed to designate both types of particles and the Greek-letter prefixes 7 and yp used to differentiate between them. ELEMENTARY PARTICLES OF PHYSICS—ANDERSON 911 Thus the term r-mesotron or r-meson designates the heavier particle and y-mesotron or p-meson designates the lighter particle. This nomenclature seemed satisfactory for a time until continued experi- mentation began to show more and more clearly the important basic differences between the two types of particles. It is beginning to be quite apparent now that the properties of these two types of particles are such that they will not naturally fall into the same classification. Thus the use of a common generic term, such as mesotron or meson, to designate both these types of particles may in the future prove to be quite inconvenient and illogical. Just what should be done with respect to nomenclature at this time is not clear, but it is a matter which should receive very serious consideration, especially in view of the apparent entry of still another new elementary particle into the fold. Another important advance that I want to mention is the recent success in producing mesotrons in the large cyclotron on the University of California campus at Berkeley. This represents the first time that it has been possible by artificial or laboratory methods to imbue a single particle of matter with an energy sufficiently high to make pos- sible the creation of mesotrons. This they have succeeded in doing in Berkeley with their beam of a-particles, or helium nuclei, which have been accelerated to an energy of 400 million electron volts. They observed the production of both the heavy and light mesotrons, and all indications are that the mesotrons thus produced are identical with those previously observed among the particles produced by the cosmic rays. Now in the design stage are other particle-accelerating machines which will yield particle energies several times the 400 million electron volts so far achieved in the Berkeley cyclotron. When these machines are in operation, working at energies up to 6 or 7 billion electron volts, we can expect to learn much more about mesotrons and the other ele- mentary particles of matter. Moreover, we must expect that a con- tinuation of research in cosmic rays will also extend our knowledge in this field, since in the cosmic rays particles are available for study whose energies are even 10 to 100,000 times greater than those to be expected from any of the accelerators that are being planned. In conclusion I should like to indicate the possible significance of these new discoveries to science and to the world at large. In this discussion I have classified physical phenomena, according to the energy associated with them, into three categories: (1) low- energy or extranuclear phenomena, (2) high-energy or nuclear phe- nomena, and (3) extremely high-energy or what we might call, for want of a better name, elementary-particle phenomena. Knowledge of the first of these, low-energy or extranuclear phenomena, has already profoundly affected the life of nearly every human being on earth. 2A ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The industrial revolution, our mechanized civilization, the shrink- ing of the world through advances in communication and transporta- tion have all come as a direct application of our knowledge of low- energy or extranuclear phenomena. Indirectly it has been responsible for the political and economic organization of the whole earth. Our present age might well be classified as an extranuclear age. Since the explosion of the atomic bomb, and the achievement of the release of nuclear energy on a large scale, it seems rather clear that we are now entering a new period in which nuclear phenomena are destined to have an important part in shaping the world, at least politically if not economically, in the very near future. Just how great will be the influence on the world of our knowledge of nuclear phenomena no one can say. It is only 50 years since our direct knowledge of the electron was not much more than a faint green glow in a glass tube—and now no one would deny that our knowledge of the properties of the electron has had an effect of profound importance in shaping our civilization. It is also only about 50 years since the world’s knowledge of nuclear phenomena consisted of nothing more than the thoughts passing through the mind of Becquerel as he pondered a darkened area on a photographic plate. At present our knowledge of all these fields is incomplete, but particularly is this true of nuclear phenomena, and most particularly true of high-energy phenomena or the phenomena of the elementary particles. So far, the world’s knowledge of the phenomena of high energies or the interactions between the elementary particles is represented by nothing more than a few printed pages in the scientific journals, by discussions among physicists, or perhaps by an occasional lecture. But we can look forward with anticipation and even excitement to the new discoveries which are surely to come as studies are carried forward of elementary particles and very high-energy processes. New phenomena of great beauty, extreme complexity, and novelty are certain to be revealed and finally to be understood. Whether our knowledge of these new phenomena will then exert a great or a small influence on the world as a whole no one can say. I believe it would be most unwise, however, in the light of the history of scientific development, to expect this influence to be small. RECENT ADVANCES IN VIRUS RESEARCH ! By WENDELL M. STANLEY Professor of Biochemistry and Director of the Virus Laboratory University of California Viruses are small infectious agents that can cause disease in man, other animals, plants and bacteria. They range in size from about 10 my, a size slightly smaller than that of certain protein molecules, in an almost continuous spectrum of sizes up to about 300 muy, a size slightly larger than that of certain accepted living organisms. A given virus can multiply and cause disease only when within the cells of certain specific living organisms. No virus has been found to repro- duce in the absence of living cells. During multiplication viruses occasionally change or mutate to form a new strain which in turn causes a new disease. Viruses were not discovered until 1892 when Iwanowski demonstrated that the causative agent of the mosaic disease of tobacco would pass through a filter that retained all known living organisms. Six years later Beijerinck proved that this agent was not an ordinary living organism and recognized it as a new type of infectious disease-producing agent—namely, a virus. The same year Loeffler and Frosch demonstrated that foot-and-mouth disease of cattle was caused by a virus. The discovery of the first virus disease of man, that of yellow fever, was made in 1901 by Reed and coworkers. Since the original discovery of the infectious, disease-producing agent known as tobacco mosaic virus, well over 300 different viruses capable of causing disease in man, animals, and plants have been discovered. Among the virus-induced diseases of man are smallpox, yellow fever, dengue fever, poliomyelitis, certain types of encephalitis, measles, mumps, influenza, virus pneumonia, and the common cold. Virus diseases of animals include hog cholera, cattle plague, foot-and- mouth disease of cattle, swamp fever of horses, equine encephalitis, rabies, fowl pox, Newcastle disease of chickens, fowl paralysis, and certain benign as well as malignant tumors of rabbits and mice. Plant virus diseases include tobacco mosaic, peach yellows, aster yellows, 1 Talk presented at the Medal Day Meeting at The Franklin Institute, October 20, 1948. Reprinted by permission from Journal of the Franklin Institute, vol. 246, No. 6, December 1948. 213 214 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 potato yellow dwarf, alfalfa mosaic, curly top of sugar beets, tomato spotted wilt, tomato bushy stunt, corn mosaic, cucumber mosaic, and sugarcane yellow stripe. Bacteriophages, which are agents capable of causing the lysis of bacteria, are now regarded as viruses. The viruses have been separated as a special group of infectious, disease-producing agents by means of several general properties, no one of which is, however, exclusively characteristic of viruses. Never- theless, no great amount of difficulty has been encountered in the segregation of the virus group. Viruses are characterized by their small size, by their ability to reproduce or multiply when within the living cells of a given host, by their ability to change or mutate during multiplication, and by their inability to reproduce or grow on artificial media or in the absence of specific living cells. The sole means of recognizing the existence of a virus is provided by the multiplication of the virus which is, of course, usually accompanied by manifesta- tions of disease. Viruses spread from diseased to normal susceptible hosts by different methods. Some are transferred by direct contact, as when a diseased leaf is caused to rub against a healthy leaf by a gust of wind, or when a normal person or animal comes into direct contact with a diseased person or animal. Such viruses can usually be spread by indirect contact through the medium of nonspecific animate or inanimate objects. Some viruses cannot be transferred by direct contact, but require an intermediate host such as a mosquito, louse, or leafhopper. Im some cases a highly specific intermediate host is necessary, and a more or less definite period of incubation within this host may be required before the virus can be transmitted. Because properties such as reproduction and mutation have long been considered characteristic of living entities, viruses were, for many years, regarded as living organisms somewhat smaller than ordinary bacteria. However, the isolation in 1935 of tobacco mosaic virus in the form of a crystalline nucleoprotein of unusually high molecular weight and the subsequent isolation of still other viruses in the form of high molecular weight nucleoproteins, some of which were also crystallizable, cast doubt upon the validity of classifying all viruses as organisms. With the exception of virus activity, the properties of some of the smaller viruses are quite similar to the properties of ordinary protein molecules, whereas at the other extreme with respect to size, the properties of the viruses are more nearly like those of accepted living organisms. The viruses, therefore, serve as a bridge between the molecules of the chemist and the organisms of the bacteriologist, and provide us with new reasons for considering that life, as we know it, owes its existence to structure, to a specific state of matter, and that the vital phenomenon does not occur spon- taneously, but is possessed in varying degrees by all matter. It is ADVANCES IN VIRUS RESEARCH—STANLEY 915 obvious that a sharp line dividing living from nonliving things can- not be drawn and this fact serves to add fuel for discussion of the age-old question “What is life?” Attempts to learn something about the nature of viruses through studies on their general properties began with Beijerinck’s work in 1898 and were continued in different laboratories for over 30 years without too much success. Although Beijerinck and Allard made important contributions, perhaps the most significant work was that of Vinson and Petre during the years from 1927 to 1931 when they showed that tobacco mosaic virus could be subjected to several kinds of chemical manipulations without loss of virus activity. Never- theless, in 1932 the true nature of viruses was a complete mystery. It was not known whether they were inorganic, carbohydrate, hydro- carbon, lipid, protein, or organismal in nature. It became necessary, therefore, to conduct experiments which would yield information of a definite nature. Tobacco mosaic virus was selected for these initial experiments because it appeared to provide several unusual ad- vantages. Large amounts of highly infectious starting material were readily available and the virus was known to be unusually stable. Furthermore, it was possible to titrate or measure the amount of this virus in a preparation with ease and rapidity and with great accuracy. During the course of a wide variety of early exploratory experiments, it was found that the enzyme pepsin inactivated tobacco mosaic virus only under conditions under which pepsin is active as a proteolytic agent. It was concluded that tobacco mosaic virus was a protein or very closely associated with a protein which could be hydrolyzed by pepsin. With this as a lead, efforts were made to concentrate and purify tobacco mosaic virus by means of the methods previously employed in work with proteins. Soon, by means of a combination of procedures involving salting-out, isoelectric precipita- tion and adsorption on and elution from an inert material, a crystal- line material was obtained which possessed the properties of tobacco mosaic virus. This crystalline material was found to be a nucleo- protein with rod-shaped molecules or particles about 280 by 15my in size and with a molecular weight of about 40,000,000. Early skepticism that a virus could exist in the form of a crystallizable nucleoprotein has largely disappeared, chiefly because the results of a vast amount of experimental work have indicated that the virus activity is a specific property of the rod-shaped nucleoprotein. Tobacco mosaic virus exists in the form of many strains which appear to have arisen by a process similar to that of mutation in higher organisms. Several of these strains have been obtained in purified form by means of differential centrifugation. Purified prep- arations obtained from plants diseased with different strains of 866591—50-——15 216 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 tobacco mosaic virus were found to possess properties quite similar to, yet in every case distinctive from, those of purified preparations of the ordinary strain. Spectacular progress has been made in the establish- ment of the nature of the chemical changes which accompany the mutation of tobacco mosaic virus. The amino acid composition of purified preparations of eight strains of tobacco mosaic virus and of two types of influenza virus has been determined. The results ob- tained with the strains of tobacco mosaic virus indicate that the mutation of a virus can be accompanied by the elimination of one or more amino acids from the virus structure, by the introduction of one or more new amino acids into the virus structure, or by a change in the concentration of one or more amino acids present in the virus structure. This work has great significance for it has provided the first informa- tion regarding the nature of the structural changes which accompany mutation. Extension of this work may reveal the exact nature of the chemical differences between virulent and avirulent virus strains, and provide important information regarding the mutation process in higher organisms. Attempts have been made to change the structure of tobacco mosaic virus by means of known chemical reactions in vitro in an effort to se- cure chemically modified active virus. Although several types of chem- ical derivatives of this virus were produced and were found to possess full virus activity, the inoculation of such virus derivatives to normal Turkish tobacco plants always resulted in the production of ordinary tobacco mosaic virus. ‘The results indicated that the chemical deriva- tives were converted to ordinary virus following their introduction into the cells of the plant, or, more probably, that the infecting molecules may not necessarily function as exact patterns for reproduction. De- spite these results it still appears that it may be possible to make changes in vitro similar to those which occur in nature, and thus secure a heritable chemical modification. Obviously this is a field in which important new results can be anticipated. Following the isolation of tobacco mosaic virus in the form of a crystalline nucleoprotein having individual molecules or particles about 15 by 280 my in size, studies were undertaken in several laboratories to determine if other viruses could be obtained in purified form, mainly by techniques involving high-speed centrifugation. Some of these puri- fied viruses are crystallizable nucleoproteins having either rodlike or spherical particles. Some are nucleoproteins which have, as yet, not been crystallized. Others are large particles consisting of nucleo- protein, lipid, and carbohydrate, and possessing, in some cases, a degree of morphological differentiation which resembles that of organ- isms. Still other viruses have, as yet, defied isolation and purification, possibly, in some cases, because of extreme instability. The viruses ADVANCES IN VIRUS RESEARCH—STANLEY Pail which have been purified possess varied shapes and form an almost continuous spectrum of sizes. The smaller rod or spherically shaped viruses appear to be simple nucleoproteins, some of which can be obtained in crystalline form. These appear to have chemical and physical properties which, neglecting virus activity, would tend to place them in the molecular world. The larger viruses have a com- position and properties which are characteristic, not of molecules, but of organisms. The viruses have certainly provided a link between the molecules of the chemist and the organisms of the biologist. Yet there is no place at which a line can be drawn dividing the molecules from the organisms. The viruses appear to form a continuous series with respect to structure, ranging from the smaller viruses, which are simple nucleo- proteins with many properties similar to those of ordinary molecules, on through viruses with a gradually increasing complexity of structure, to the larger viruses, which, with respect to structure and properties, are similar in many respects to organisms. However, it must be re- membered that the properties of only a relatively few purified viruses have been determined. In view of the possibility that these represent the more stable and more easily purified viruses, one cannot be certain that a true picture of the chemical and physical properties of viruses as a whole has been obtained as yet. Information regarding the mode of reproduction of viruses is needed most urgently. At present it is not known whether viruses reproduce by fission or by means of some new process. The solution of this puzzle would certainly represent a most important and significant advance, for the basic reactions character- istic of virus reproduction may well represent the fundamental process which characterizes all living things. GROUND-WATER INVESTIGATIONS IN THE UNITED STATES! By A. N. Sayre Geologist in Charge, Ground Water Branch Water Resources Division, U. S. Geological Survey Before discussing ground-water investigations in the United States I should like to outline briefly the broad problems of water supply, water control, and conservation, in the solution of which ground- water investigations play an important part. For mere existence, a man requires only about 2 quarts of water per day. However, even in simple pastoral or agricultural settings, the biological necessity represents only a part of the total needs for water, and in our own complex industrial and agricultural economy great quantities of water are needed for a multitude of purposes. Water is needed for sanitation, for washing clothes, for facilitating sewage disposal, for scrubbing floors, and for processing foods. It is needed for fire protection, for generating power, and for industrial processes, for irrigation, for air conditioning, and even for producing atomic energy. The task of providing water at the right time and place is a serious problem which fully occupies the attention of thousands of engineers and chemists and a smaller number of geolo- gists. Intimately associated with the water-supply problem is the problem of controlling floods to minimize the erosion of our soils and to conserve floodwater for beneficial uses. For obvious reasons, many of our agricultural, industrial, and urban developments have taken place along waterways where they are vulnerable to the ravages of floods which appear to become more costly almost year by year. There is good reason to believe that the demand for water supply will continue to increase and that the demand for control and conservation of floodwaters will also increase. Projects for accomplishing these ends will become more expensive and their planning and design will require greater knowledge of our water resources and of the basic geologic and hydrologic factors affecting them. 1 Presented at the joint meeting of the Society of Economic Geologists and Geological Society of America, Ottawa, Canada, December 1947. Reprinted by permission from Economic Geology, vol. 43, No.7, Novem- ber 1948. 219 220 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Especially during the past century, the use of water has increased at an amazing rate. In 1850 only 83 cities in the United States had public water supplies, and only a small proportion of the homes in these cities had water piped directly from the city mains. By 1939 there were 12,760 ? municipal waterworks and thousands of industries had private supplies from wells or surface-water sources. As the number of waterworks increased, the uses of water and the per- capita consumption also increased. The quantities of water used for a few purposes are given below. Flush toilets, bathing and laundry, street cleaning, and fire protection require an average of about 40 to 75 gallons per day per capita.* Processing a ton of steel in highly finished form requires about 65,000 gallons* of water; making a gallon of gasoline takes 7 to 10 gallons® of water. Vast quantities of water are used for air conditioning and for making paper, explosives, coke, textiles, and a host of other products. Thus, a large city, such as Chicago, with numerous industries may have a per-capita water consumption as high as 250 to 300 gallons per day. Only a few thousand acres in the West was irrigated in 1850, but 21 million acres ® was irrigated in 1939, and many additional irrigation projects are under construction. An acre of cotton uses about 2.57 acre-feet, or 800,000 gallons, of water during the growing season; an acre of alfalfa requires about 4 acre-feet of water; irrigation of truck gardens, fruits, sugarcane, rice, and other crops also requires large amounts of water. In eastern United States, supplemental irrigation is increasing because the application of a relatively small amount of water when it is needed by crops may double or triple the yield of the land. The greatly increased use of water has, in many places, almost fully utilized the readily available water supplies, drawn ground-water levels dangerously low, caused sea water to enter streams and ground- water reservoirs in coastal areas, and permitted oil-well brines or factory wastes to pollute many of our ground-water reservoirs and streams. Thus, the development of additional water supplies for new projects or industries has become increasingly difficult and costly. Nevertheless, it would be a mistake to infer that our water supplies are approaching exhaustion. Actually, much can be done to conserve and thereby increase the total amount of water available for beneficial use. For example, in many places spacing pumped wells over wider 2 Engineering News-Record, vol. 123, p. 414, 1939. 3 Turneaure, F. E., and Russell, H. L., Public water supplies, 4th ed., p. 19. John Wiley & Sons, New York, 1940. «Lloyd, Kenneth M., Industry and water supply in Ohio. Ohio State Univ. Exper. Stat. News, p. 31, April 1946. ‘ Jordan, Harry E., Industrial requirements of water. Amer. Water Works Assoc. Journ., vol. 38, pp. 65-68, 1946. 6 Census Bureau. ’ National Resources Committee, Regional Planning, pt. 6, p. 91, 1938. GROUND-WATER INVESTIGATIONS—-SAYRE 221 areas would prevent excessive lowering of the ground-water levels. Artificial recharge of ground-water reservoirs by spreading flood- waters and by other means has been successful in several areas. Abatement of pollution in streams and ground-water reservoirs, retention of floodwaters in reservoirs for later use, control of reservoir stages by forecasting normal and flood flows of streams, control of silt and sedimentation, and other measures are being carried out to increase the supply available for perennial beneficial use. The continued growth and prosperity of the Nation will depend to a large degree upon the success with which these problems are attacked and solved. Unlike most mineral resources, water is not exhaustible, in the strict sense, because it is replenished from time to time by precipitation. A surface reservoir may be dangerously low and be refilled in the nick of time by heavy rains. Heavy pumping may cause ground-water levels to decline progressively until pumping is no longer economically feasible, but when pumping is temporarily or permanently reduced the water levels usually recover. Likewise, a period of heavy rainfall following a drought period may induce recharge sufficient to restore water levels essentially to predrought levels. Only a part of the water taken from streams or pumped from wells is actually consumed. In many manufacturing processes water is merely a washing agent and is essentially unreduced in volume by its use. Water used in boilers or in quenching hot metal is partly evaporated, but the remainder is discharged or re-used. Water used in irrigation is partly evaporated and partly transpired by plants, but there is always an excess which is discharged and which carries away undesirable salts. The excess water from all these uses returns to the stream or to the ground altered by the concentration of minerals contained in it, or by the addition of dissolved constituents, or of color, or sediment, or simply by the addition of heat. It may be re-used for the same or other purposes with or without the addition of new water. For example, the water of the Pecos River, in Texas and New Mexico, is used and re-used for irrigation and domestic supply seven or eight times between its source and Girvin, Tex. Al- though water is added from tributary areas along its course, each time the water from the river is used the mineral concentration increases, and a few miles above Girvin it is so highly mineralized that even the most resistant crops are unable to survive its application. Even so, it may still have potential use, because water also possesses the energy of position and in its journey from the mountains to the sea it may be used many times over for generating hydroelectric power. Another characteristic peculiar to water results largely from the vagaries of precipitation. Many places are faced successively with Dae, ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 water shortages and destructive floods. Although enormous sums have been spent on flood control, complete protection from floods is difficult and often prohibitively costly, and in only a few places has it been accomplished. Whatever conclusions are reached with regard to the economics of flood control by storage reservoirs, from the standpoint of conservation of water supply it is advantageous to salvage as much of the floodwater as possible. In certain areas where water supply is now inadequate, as it is in many places west of the 100th meridian, much of the water from precipitation escapes to the sea during floods. In Los Angeles County, Calif., much of the floodwater is retained in surface reservoirs from which it is later discharged into specially prepared recharge basins and seeps into the underground aquifers. Some of the flood- water is caught in seepage reservoirs designed to promote infiltration into the ground-water basins, and some of it escapes to the sea, but essentially all the floodwater in the Los Angeles area will be put to beneficial use when the projects now under consideration are com- pleted. Plans for similar flood control in other parts of the country are in various stages of execution. Ground water and surface water are so intimately related that for proper solution of the over-all problems of water supply, control, and conservation it is now necessary to have all the facts regarding both. Precipitation, which is the source of both, is partly lost, largely through evaporation. Especially during the growing season, a large part enters the ground and is transpired by plants. The remainder per- colates downward below the plant roots to become ground water, or runs off directly as surface flow. The ground water, returning to the surface as seeps or springs, provides the base flow of the streams which prevails through periods of low precipitation, On the other hand, especially in the West, many streams lose water by seepage in certain stretches and thus recharge the ground-water reservoirs. Many of the basic ground-water investigations in the United States are carried on cooperatively by the United States Geological Survey and State or local agencies, including State geological surveys, State engineers, counties, and municipalities. In Illinois ground-water investigations are made by the State Geological Survey and the State Water Survey; and in Missouri investigations are made by the State Geological Survey. In California a large staff of engineers in the State Division of Water Resources for many years has been investi- gating overdraft of ground-water supplies. Recently arrangements were made whereby the United States Geological Survey, in addition to its investigations in Los Angeles, Orange, and Santa Barbara Counties, will assist the Division of Water Resources in the geological phases of a State-wide inventory of the water resources of California. GROUND-WATER INVESTIGATIONS—SAYRE 223 Various other Federal and State agencies are obtaining some data on ground water in connection with special phases of their work. The ground-water investigations of the United States Geological Survey began more than 50 years ago. At that time ground-water supplies were little developed. Consequently, most of the early field investigations were of the exploratory type. Laboratory and field studies by King, Slichter, and later by Meinzer outlined the broad principles of ground-water occurrence and movement. However, before the deep-well turbine pump was developed in the early part of this century, use of ground water in large quantities was limited to areas of springs or artesian flow, or to areas where the water level was within reach of suction pumps. After the turbine pump was intro- duced, it became possible to pump economically even where water levels are deep; power costs also decreased, and well drilling and finish- ing methods were improved to increase the efficiency of wells. Because of these advances, ground water came to be used in ever-increasing quantities, first in areas where surface water was not readily available, and later in areas where ground-water supplies were more economical because they obviated the long pipe lines, collection works, and costly treating plants needed for surface-water supplies. ‘The first great expansion of ground-water supplies was made largely without technical guidance. Because of their immense storage capacity, the ground- water reservoirs were regarded as inexhaustible, and in many places development was well advanced before progressively declining water levels brought about the realization that ground-water reservoirs may be depleted. As a result, demands for detailed ground-water studies steadily increased. These studies are thorough, systematic investiga- tions which include areal geologic mapping; subsurface studies occa- sionally augmented by geophysical surveys and test drilling to deter- mine the structure, thickness, and the sequence of water-bearing and non-water-bearing beds; collecting data on fluctuations of water levels in relation to precipitation and to pumpage; determining the recharge and perennial yield; inventories of ground-water withdrawal; test pumping to determine coefficients of permeability, transmissibility, and storage; determining the quality and temperature relationships, and the relations between surface and ground water. The work does not include supervision, construction, or control of water supplies. The United States Geological Survey is now making cooperative ground-water investigations in 42 States and in Alaska, Hawaii, the Virgin Islands, and Puerto Rico. In general, the State or local authorities are most familiar with the needs in their States, and they are largely responsible for designating the areas in which investigations are to be made. Most of the ground-water staff of the United States Geological Survey now have headquarters in field offices, of which there are about 40. B24. ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 In addition to the cooperative investigations with States and local agencies, certain investigations primarily of Federal interest are being carried out with use of Federal funds only. For example, an extensive program of ground-water investigations is being conducted in the Missouri River Basin to provide basic information needed for planning and constructing the various water projects that are authorized or planned. This information will include determination of existing ground-water conditions and probable effect of irrigation on ground- water levels, especially with reference to waterlogging and drainage, canal locations with reference to seepage, areas where irrigation with ground water is feasible, the location of potential sources for farmstead and municipal water supplies, and so on. Other investigations, such as that recently started in the Central Valley of California, will provide basic information needed in the project to use excess floodwaters for recharging the heavily pumped ground-water reservoirs in the San Joaquin portion of the valley. Several projects also are under way in connection with defense plans. A large program of measurement of ground-water levels in observa- tion wells has long been an integral part of the cooperative program. Recently a small Federal fund was provided for extending this pro- eram, analyzing the data, and determining the current status of our ground-water supplies on a Nation-wide scale. The program includes the investigation of the possibilities of forecasting low-stage stream flows from fluctuations of ground-water levels, enabling more efficient control of reservoir and river stages in the operation of hydroelectric plants and permitting considerable economies in the generation of power. Conversely, base flow is an index of ground-water storage. As the ability of the ground-water reservoirs to absorb water ma- terially affects the runoff to be expected from rainfall, the analysis of data on ground-water levels should aid in flood forecasting. In addition to and as part of the above program, several research projects are under way, including the study of movement of water through soils and water-bearing materials; infiltration from streams and its effect on the temperature and quality of the ground water; the occurrence of ground water in fractures and solutional openings in impermeable rocks, such as limestone, tightly cemented sandstone, granite, etc., and the improvement of our techniques for locating producing wells in such rocks; earth subsidence resulting from the withdrawal of ground water, and related problems connected with elasticity and compressibility of artesian aquifers; methods of analyz- ing results of pumping tests to determine coefficients of transmis- sibility and storage; and a number of other projects. With respect to the probable future of ground-water investigations, it should be pointed out that ground-water development was expand- GROUND-WATER INVESTIGATIONS—SAYRE 225 ing rapidly in the middle thirties. In 1935 the total use of ground water in the United States amounted to about 10 billion gallons a day. The development was greatly accelerated by the needs of the war, so that by 1945 the total pumpage had nearly doubled.2 Although the war ended more than 2 years ago there has been no sign of a decrease in the use of ground water. In fact, both surface water and ground water are now being used in greater quantities than ever before. As the use of water approaches ever more closely the limits of the available supply, it is believed that water-resources investiga- tions will be needed with ever-increasing urgency because water constitutes the prime factor in the continued development of the Nation’s industrial and agricultural economy. § Guyton, W. F., Industrial use of ground water in the United States. Abstract, Journ. Washington Acad. Sci., vol. 39, No. 3, pp. 105-106, Mar. 15, 1949. (To be published in Proc. Geol. Soc. Amer.) es ‘ gations ban tat ipiecepyiieaseeis dur ai : once ange. ‘ aaly Ampust ra yey & ioe Gey efida-letiie Py aia oa: Aes #1) dl orang heh hae, | aa ane ON, ue fra Bia neva, Tih yt 10% 0, ast) ssa, ai a + hosts n Sousa a har wiih, af te PY, A of nid thy guys $ rhs ia oldiol ohh to tbonit,od, leg exit) 9 SFE finger thrton ay, oe ‘ies sank ARG, out “olga, pO CO lark sthen a has eo tay at) Sif wi j 4 “| i ev iccuvans AeecT Oper) ay, i iy yibitan? eifal Ai nent uy oat aah iy Hitt pF mide Haid i) ni Fat yhhgabh lt tae = 45 Vols CMTC. 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Often it is included with the physical sciences, like chemistry and physics. Yet it is also grouped as correctly with the biological sciences, along with plant physiology and bacteriology. Then again, soil science is appropriately grouped with geology and geography as an earth science. Actually soil science uses the prin- ciples and methods of all three of these groups, and a soil scientist who uses the principles of only one or two of the groups, to the exclu- sion of the others, can only have small principles that have little prediction value and soon bog down in contradiction. In addition to those already mentioned, there are important principles and methods peculiar to soil science itself that do not belong to any other science. In application, the principles of soil science must be inti- mately related to those of the social sciences. It needs to be emphasized that the experimental method and the method of scientific correlation are essential in both fundamental soil science and in the application of its principles to practical prob- lems in farming, gardening, and forestry. Soils must be studied in relation to one another and to the whole environment, both natural and cultural, to understand their forma- tion and the influence of the individual factors of climate, vegetation, parent rock, relief, and time. How any one of these factors operates depends upon the others. The significance of any one soil charac- teristic depends upon the others. Any soil is a combination of characteristics, produced by a combination of factors, each of which influences the functioning of the others. Experiments are needed, both natural and artificial, to learn how individual soils behave and how they respond to treatment. These must be specifically related to individual kinds of soil, however, if the results are to be used in developing principles or as the bases for practical predictions. Thus the experimental methods and the methods of scientific correlation are intimately interwoven in produc- tive research in soil science. SOIL AND LANDSCAPE Let us look briefly at the implications of this concept of soils. They are natural bodies, each with its own unique morphology; they are dynamic bodies, developing with the natural landscape itself; they ac- curately reflect, at any moment, the combined or synthetic influence of the living matter and climate, acting upon the parent rock through processes conditioned by relief, over a period of time; they are dis- tributed over the earth according to orderly discoverable and definable geographic principles. The geological process of mountain building, rock formation, and landscape evolution from which the parent materials of soils originate, MODERN SOIL SCIENCE—KELLOGG 25K are still going on along with soil formation. The natural erosion of the uplands gradually removes a little of the surface bit by bit while the soil film settles down, and fresh minerals are added to the soil from beneath. With the warping of the landscape these processes are accelerated or retarded. At several of the Soil Conservation Experiment Stations of the United States Department of Agriculture, rates of erosion were deter- mined under permanent vegetation (15).2 Under the natural forest Exchangeable Cations (M.E,/100gms.) Exchangeable Cations (M.E./100gms.) pH 10 20 30 40 pH 5.1 4.8 49 4.4 Si 6.2 $.6 B. From granite a ed, Percentage of Clay A. From diabase Fiaurs 2.—A comparison of the clay content and exchangeable bases from two soils developed in similar environments, except for the difference in parent rock. The fundamentally important differences in clay content and in ex- changeable bases are obvious between the soils and among horizons within each soil (9). cover of the Cecil soil of the Piedmont, erosion proceeds at a rate of about 1 foot in 10,000 years. This value was determined on a 10- percent slope. Yet on a 14-percent slope of the Muskingum silt loam of Ohio, considerably over 200,000 years would be required to remove 1 foot by erosion under the forest cover. Under a well-established grass cover, normal erosion proceeds slowly on the dark-colored soils developed under tall prairie grasses. On the Marshall silt loam near the Nebraska-Iowa line, with 9-per- cent slope, nearly 14,000 years would be required to remove 1 foot under bluegrass. On a black soil of east Texas, Austin clay with a slope of 4 percent, the figure is nearly 900,000 years. 3Numbers in parentheses refer to literature cited at the end of this article. 866591—50-——_16 Zaz ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 These values probably do not represent the real extremes, but rather relate to normal erosion. Some soils erode much more rapidly under clean cultivation with bad effects upon soil productivity. Through proper cropping systems and soil management practices, erosion of soil under use should be kept somewhere near the normal rate. Organic Organic Motter(Z) pH SOW ue e100 0) 50 100. ~— PH Matter() eth lie 3 posse TMT | Depth Pe l ae 55 6.2 Serere, iO ¥ 1 oso SEMIN ETM ss > (] ZOmennS'5 seeeene yA the ise a on a < Qj } Ie os Femme aN O 20" — Rey 6 4 lo 82 SN eee Tl, 50:7. nt 64 se : Sees ieee Gee 7.8 08 HO. 7:6 cle? oO res : RRO ! A. 40” PGR CERI Rt iat is) A. Miami Stit Loam- Undulating to Rolling B. Bethel Silt Loam- Flat Ficur& 3.—A comparison of the mechanical composition of two soils developed from similar parent materials and in similar environments, except for relief. On soils of undulating to gently rolling relief there is continually a small amount of erosion in the natural landscape. As the surface is thus slowly eroded, each soil horizon works down into the one beneath. Over a period of several thousand years the whole soil profile, while remaining at the same length, may sink into the landscape a foot or more. Thus the soil is kept constantly renewed with new minerals. This is illustrated by the Miami silt loam at the left. The soil at the right is developed from similar material on flat upland where there is little or no erosion in the natural landscape. The leached material accumulates at the surface, and clay formation and the accu- mulation of clay are accentuated in the middle portion of the profile, with the development of a claypan. Whereas the Miami silt loam is a well-drained soil, pervious to roots and water, the Bethel is an imperfectly drained soil. During wet periods excess water is held up by the claypan which is also impenetrahle to mast roots. Kinds of crops and soil-management practices for optimum production are quite different indeed, even though the soils lie side by side on the same farm (2). Percolating water gradually dissolves the minerals in the soil and the rock beneath. Clarke has said that this process alone reduces the surface of the United States on an average of about a foot in 30,000 years (6). From a study of streams, Dale and Stabler (7) estimated MODERN SOIL SCIENCE—KELLOGG 233 several years ago that the average rate of denudation for the United States as a whole was about 1 foot in 8,760 years, with suspended matter accounting for about 65 percent and dissolved matter 35 per- cent. Solution progresses much more rapidly than this in areas of soft limestone and high rainfall, and, of course, the process is almost infinitely slow on the hard rocks of steep slopes. Other landscapes receive part of the erosion products and part of the solution products from the upland. It is probable that a third of the population of the world get their major food supply from alluvial souls recently rejuvenated by additions of fresh rock minerals to their surface. The Nile is a famous example. In flood stage the water of the Nile contains over 1,000 parts per million of suspended matter relatively rich in phosphorus, potassium, nitrogen, and other plant nutrients. Part of this covers the soil in the flood plain and part of it moves out into the sea. Barrell (1) estimated that the Nile Delta alone contains the equivalent of nearly 12,000 cubic miles of rock, to say nothing of the soluble material contributed to the sea water. According to Barrell’s figures the rock material in the delta of the Niger River is equivalent to a wedge-shaped mountain range some 18 miles wide at the base, 3 miles high at the top, and 1,000 miles long. This gives some idea of the enormous movement of surface soil material as a natural process. In addition, in the Tropics especially, volcanoes often shower the landscape with fresh rock or ash. mates ‘ LE pee Af: Le Z Ses ELE row ae SA The areas of each great soil group shown on the map include areas of other groups too small to be shown separately Especially are there small areas of the azonal and intrazonal §roups included in the areas of zonal groups AZONAL Soils without well-developed soi! characteristics. (Many areas of these soils are included with other groups on the map.) LITHOSOLS AND SHALLOW SOILS oe ll Fo O46 F O)) a Shallow soils consisting largely of an imperfectly weathered mass of rock fragments, largely but ee | not exclusively on steep slopes. Resear. RON BF SANDS (ORY) Very sandy soils. ALLUVIAL SOILS Soils developing from recently deposited alluvium that have hadlittle or no modification by pro = cesses of soil formation. EK 866591—50 (Face page 236) Thad. VTE Yiya® ' “ate a? Hind “Too5 _ S a Bypoe runtesid to We Hind - #100) abbateenry td eaten etree ale Ss: paca eal Sean kee: <7 a. etn AR ecmeiieaall oa lie LEG » ontardignns. shay tanita 'o equrg tev) Fenpfeipen Fh alae Ti YO sone tes prvlenemah act yetael ‘ey byes are Nheere eur gem ett ro peo 24) > (dabviow ne ele pew ate to shoe dares! beroind yi.t —_— soto 39 m 7 2102 Anime» HeIOo aR 5 oe dey r Dun 2 hae Memo Geo ber gra ‘ tea uve bo baw eneiey unsinse bamud e604 matteo \e chee berioeg! award eee ieGupers a cb cath: abr te ef - aipay “oooge balesrot N Wros 691. cy al on t ; oe . ‘ ’ pa trem wedt f pees WO tt = pg BOR. 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Les ign +t pte ts ” seta pa 44 MODERN SOIL SCIENCE—KELLOGG De Th causes of soil characteristics, because of failure to see the common characteristics of unlike soils found in the same ecological region. In a broader study one might reach altogether different conclusions. Suppose, for example, that one studies in detail the upland soils on smooth slopes from limestone and sandstone in West Virginia in con- trast to those on smooth slopes from limestone and sandstone in western North Dakota and in southern Arizona. Here again, striking differences are apparent, but the most important of these relate to differences in climate and vegetation; geology would seem relatively unimportant. Thus by grouping the local soil types into higher categories, the broad soil groups that dominate the landscapes of great regions are compiled. When one considers individual practices in the garden, on the farm, or in the forest, it is the local landscape and the local soil types that are important. When one considers the great movements of population, the potentialities of nations, and the historical trends of peoples, the significant factors are the great soil groups. Thus the gardener sees soil characteristics with a different emphasis than does the geographer, but soil science has much to contribute to both. PRIMITIVE SOCIETIES ON THE SOIL The soil supports plants, animals, and man himself. Primitive man must have been as much the helpless product of his environment as were the wild animals. He lived close to the soil and was a food gatherer. He took the plants and animals, including fish, that were available in his own landscape. Most of these were eaten with little change by cooking, storage, or refining. No doubt it was a risky busi- ness. Families might fail to get food because of drought, deep snows, wars, or other calamities, and starve. However, when things went well, although they might live on a few foods for a time, during the year there was usually a variety. Then too, primitive man ate vigor- ously. He ate whole foods—skins, hulls, seeds, and other parts that are now thrown away. It is only recently that modern man has attempted to create a balanced diet. Early man received or failed to receive his proteins, minerals, and vitamins, unconsciously. Of course, no scientist was available to study our savage ancestors at the time, but many studies have been made of relatively primitive peoples and of the physical degeneration caused by the substitution for native whole foods, like cereal grains, milk, cheese, fruits, and meat, of refined sugar, white flour, and similar products of more ‘‘advanced”’ societies (12). Thus native peoples may become degenerated, or even extinct, after contact with Western Europeans. ‘There are reasons to suggest that people, like plants and animals, over a long period come to an adjustment with their food supply (11). 938 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Marett held that people having food deficient in some essential ele- ment, say calcium, phosphorus, or iodine, gradually develop the ability to conserve this element. That is, in the evolutionary process the ability to get along satisfactorily with only a little of some element would have survival value. Such people would be most likely to have children able to thrive and grow strong. But in the process, size, skin color, and other features, even the psychological characteristics and social traits, are altered. Marett believed that differences in food composition, most of which were closely related to the local environ- ment before the modern period, had a great deal to do with the origin of races, with the physical and social differences among the different peoples of the earth. Marett argued, for example, that in regions with acid soils deficient in calcium, people of small size would be favored. The physiological strain of lactation upon females would be much greater in humid regions where soils are generally leached and acid and the foods de- ficient in calcium and phosphorus, than in arid regions where soils usually contain abundant lime (calcium). Thus on acid soils where food is deficient in calcium the bodily strain would lead to adjustments for economizing lime through decreased size, especially of bones. He suggested that the operation of these forces may have been important in the development of fine bones among modern people as contrasted to our more coarse-boned ancestors. Such changes are very gradual, and are not marked until after long-living in a particular landscape. MAN AS A CULTIVATOR As civilization developed, man became a cultivator. He began to direct the course of nature toward his own ends, and ceased to be simply a food gatherer dependent only upon the natural bounty of the landscape. He ceased to be primarily a thief, and became a grower, 2 homemaker, a planner, and a conservationist in the only sense the term has any social meaning. As he gained in experience he learned to satisfy himself more easily; in fact, some people in the society could cease to be food gatherers. Social structures rose with the evolution of trades and professions. As the efficiency of food production increased, more and more people could be released from food gathering to develop the arts and sciences, to make the other things man needed for his health and comfort, and, unfortunately, to make war. From the dawn of history to the rise of modern science the accumu- lation of learning about agriculture was a terribly slow process. Experience, which was passed down from father to son over the genera- tions, was the only guide. Only a few departures were made, because there was no substitute for such experience. Further, there was little MODERN SOIL SCIENCE—KELLOGG 239 realization that experience on one soil, in one landscape, could not be relied upon where another soil was involved. Migrations were often disastrous for this reason. Then, too, world history records changes in the landscape under the very feet of the farmer, like the slow spread of the Sahara Desert as it gradually expands to its former position, following the moist period of glacial times (8). Unfortunately, the early scientists of Greece and Rome reached little into the problems of agriculture. With a few conspicuous excep- tions, the philosophers of that day accepted farming as the job of slaves, beneath the dignity of trained scholarship. THE GREAT DISCOVERIES The tempo of man’s struggle with his environment completely changed with two great forces: the rise of modern science and the great discoveries. The most important fact of Western culture was the opening of new land in the world. The forces leading to the pessi- mism of Malthus were already destroyed before his famous essay on population had been printed. Science began to increase productive efficiency. Western Europeans found new homes in the landscapes of the Americas. Europe had been bound by an aristocracy based upon land. Al- though many came to the new world to seek gold and adventure, most people came to find land and to build homes on the only security they knew. Gradually the east coasts of the new world filled up. In the beginning people were confined to land near the sea and to navi- gable waters, as they had been in the centuries before. But modern science came to the aid of discovery. The European colonists pushed into the interior, especially in North America. Railroads had made possible the exploitation of interiors of continents, of the great areas of black soils. Except for a few isolated spots, these soils were scarcely used by civilized folks at the time of the Treaty of Westphalia when modern nationalism had its birth. During the nineteenth century the black soils (Chernozem and Prairie soils) and the brown soils (Chestnut and Brown soils) in North America were occupied. The frustrated, the persecuted, the seekers of new opportunity had a place to go, and it was a good place with good soil. For over 200 years a man and woman could carve themselves out a farm home on the colo- nial frontier, in the Ohio Valley, on the great prairies of Illinois and Iowa, and finally on the Great Plains to the west. Grassland needed only cultivation, once transportation was established. In addition to the fine soils, there were other free resources, the forests and minerals. There was land in North America, Central America, South America, New Zealand, and Australia. Following the great discoveries Europeans found opportunities throughout the 240 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 world. This progress is credited by many people to western civiliza- tion, but it would have been a poor civilization indeed that could not have succeeded with these riches. With such abundant resources for its citizens, it is hard to imagine how any American government could possibly have failed. At the present time, however, people must make a go of things where they are, or else move into areas where a great deal of careful planning is necessary for successful agriculture or industry. There is no more Chemical Composition (%) Chemical Composition (%) CaO MgO oO 50 100 100 CaO MgO 0 50 Ol 02 Rosie _| Depth RQIYGY By 04 2.0 O te NW ae RA XO QOS oo .0 OS 15.4 0.03 35.5 —AAACDSL Xe oe O25 0: B. Conomingo Silt Loam of Maryland 0.2 Tr 100 a ET AP TE I TF LAD LT LD A 7 Eee == FIO OKD KOO COCO O Oe, RK KRG ROKER KK OL i Nes SKS SERRE SPRERS ‘ “ Nas OPS RRR SRKRKKS OKI K RK KR K POS Nee ROKK LKR KR SKLAR eataces xO eatetstctes ate = 25 PARAHIPPUS PARAHIPPUS . MIOHIPPUS 30 t 5 wy MIOHIPPUS MESOHIPPUS O 3544 O ‘ 2 16 re) MESOHIPPUS 40 | 45 | 9 a aii OROHIPPUS 50 I HYRACOTHERIUM 55 HYRACOTHERIUM Ficure 1.—The evolution of the horse in North America correlated with the absolute time scale. (Based on the work of E. D. Cope, W. D. Matthew, W. B. Scott, G. G. Simpson, R. A. Stirton, and others.) 250 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 nor even in successive species of ages as much apart as 1 million years, to believe that the minute differences present had any value as a determining factor in survival. Returning to the duration of a genus, it will be interesting to consult other groups, to see whether the figure of 5% million years found in the horses is more generally applicable. Similar values are found in other groups of Mammalia, such as the land carnivores. For these Simpson constructed a ‘‘survivorship curve’ showing that the mean duration of a genus of carnivores is 6% million years. But for the bivalve Mollusca, he obtained a mean as high as 78 million years. Just to add a few more figures: Swinnerton found 20 million years as the average lifetime of a genus of Triassic ammonites. Lingula, on the other hand, a brachiopod, has been known for its persistence since the Cambrian. This genus has lasted over 400 million years. From this evidence the terrestrial Mammalia appear to have genera of a short duration, but even this is a matter of a few million years. One might be inclined to interpret these different rates of generic evolution in terms of species steps, assuming that each lineage would pass through several evolutionary changes, each constituting a new species, before the accumulated differences justify one in calling the descendants a new genus. It is certain that this process underlies the evolution of the horse, but it is conceivable that in other groups genera may have originated without the interposition of a series of species steps. The time rate of species formation, therefore, must now be considered. Selecting again terrestrial forms of life, mainly mammals, as our first examples, because more detailed evidence is available, we find that since the end of the last glaciation, some 10,000 to 20,000 years ago, only minor subspecies have appeared. The differences are confined to body size, color, slight differences in body proportions, in the development of appendages, etc. The British race of the red deer, for instance, has evolved since Britain became separated from the Continent some 7,500 years ago. This is shown by fossil evidence, since the early postglacial specimens found in the Thames belong to the Continental race. Moreover the characters of Cervus elaphus scoticus, as the British race is called, are probably not fixed genetically, since, when the breed was transferred to a favorable environment, in New Zealand, it reverted in many respects to the Continental type. Other examples could be given of the insignificant character of Postglacial differentiation in species. ; Greater differences are observed in some, but by no means the majority, of Upper Pleistocene species, of about 50,000 to 100,000 years ago. For the marmot of that time, for instance, Wehrli found that the shape of the temporal ridges has since become stabilized. TIME IN EVOLUTION—ZEUNER 251 These ridges run into the upper posterior edge of the processus postorbitalis in a certain number of fossil specimens, while in recent Marmota marmota they have moved to the upper side of the processus. That this character has become practically fixed since the Upper Pleistocene, is shown by the following figures: TaBLeE I Marmota Inter- Primitive tyz.e mediate type (Percent) (Percent) (Percent) Wppersbleisvocenes 32. 4- =) = oe a IS 33 53 13 FES COT Gee ae a a res ee a ae Byer eye I a 98 LM, 2 Forms of this degree of variation would, in the recent fauna, be re- garded as subspecies. Going back to the Middle Pleistocene, about 250,000 years ago, the differences become more conspicuous, but they are still treated as subspecific by most taxonomists. It is only in the Lower Pleisto- cene, about 500,000 years ago, that we encounter ancestral forms on which the taxonomists agree that they must be classified as distinct species. This applies, for instance, to the European elephants, Elephas antiquus, and the mammoth, which Soergel has shown to have evolved from an early Pleistocene common ancestor, Hlephas meridionalis. If this view is correct, nearly half a million years were required to produce a new species by way of a gradual change. If it is not correct, the point of divergence lies farther back in the past and the rate of species change is longer. In the lineages of rhinoceroses, bears, and deer, similar evidence is available and there are cases in the Recent fauna which have been explained as the results of geographical isolation during the glaciations (the common crow and the hooded crow, for instance). But again there are species which have changed much less since the Lower Pleistocene, which are regarded as no more than subspecifically distinct from their Recent descendants and, therefore, must have a slower rate of species formation. In the Cromer Forest Bed (about 500,000 years ago), 14 percent of the species are regarded as “Recent,” though it is quite likely that subspecific differences will be detected by future revisers. Five hundred thousand years, then, appears to be a fast rate of species evolution in terrestrial mammals and no faster rate has, to my knowledge, yet been found in other groups of the animal king- dom. Let us now consider the question, how long have species re- mained stable or unchanged? To answer it we have to resort to ma- rine groups. Lower Miocene Mollusca of Java, for instance, are regarded as conspecific with Recent forms. They have been stable for 30 million years. Surveying fossils in general, this has to be regarded as a high figure, but it need not be a maximum. 252 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 It is conceivable that plant species behave differently. Small has analyzed several groups of plants from the chronological point of view, notably the diatoms. He came to the conclusion that the mean duration of species is measured in millions of years. Furthermore, he distinguished in the diatoms long-lived species (up to 100 million years) and short-lived species (one million years or less). In other groups he encountered shorter rates than this. These time rates are of much the same order as in the animal kingdom. But there is one interesting difference: while in animals the evidence suggests gradual change of specific characters, Small regards the changes in the diatoms (and perhaps other plants) as sudden. Summarizing, we may say that half a million years is a short life- time for a species and, so far as our admittedly incomplete evidence goes, something ike a minimum. The lifetime of genera appears to be rarely less than 5 million years. Short rates are observed in ter- restrial groups like most Mammalia, while long rates are frequent in mainly marine groups like the bivalve shells and the diatoms. There appears to be a certain amount of correlation between the rates of species evolution and the degree of changeability of the environment, slow rates being frequent in environments like the sea in which living conditions are exceedingly stable. Hitherto we have been discussing changes in single lines of descent. Let us now consider the phenomenon of phyletic splitting. Branch- ing is a frequent event in phylogenesis. An ancestral species may evolve into two divergent descendant species by the disappearance of intermediates, or the ancestral species may remain unaltered and a new type emerge as aside branch. In fact, few groups would survive without splitting, since owing to the action of internal (genetic) and external (environmental) factors, many species become extinct. If the rate of splitting equals the rate of extinction, the number of species in a systematic category, like a genus or a family, remains constant. But if the rate of splitting is greater than the rate of ex- tinction, the number of species in the group under consideration will rise increasingly steeply along an exponential curve. There are many different ways in which the rate of splitting can be plotted in relation to time. I have selected two simple methods of plotting which can be applied readily to any group from which suffi- cient fossil material has been adequately studied by a taxonomist. One is to plot the number of species existing in any period or sub- period, or smaller stratigraphical division, on the absolute time scale. The other consists of doing the same for the newly appearing species only. In the first, the surviving species modify the picture. The second is a truer presentation of the rate of splitting, but the material is often not complete enough for its application. TIME IN EVOLUTION—ZEUNER 253 Now let us consider a few examples. Figure 2 shows the genus Salenia. These sea urchins start in the Lower Cretaceous. The number of species rises rapidly to a climax in the Upper Cretaceous (“increase phase’”’). Cases like this one, of a rapid, almost sudden, blossoming-out of a group were called ‘explosive evolution” by Schin- dewolf. We may borrow the term and call the phase of rapid increase the explosive phase. In Salenia it is followed by a catastrophic drop followed by a slow ‘‘decline phase” leading to almost complete extinction. Of the cause of this sudden drop we are completely igno- rant; it is an unusual feature, as will be seen when other diagrams are considered. As regards the increase phase, we learn that in Salenia it lasted for about 40 million years. Figure 3 shows the Brachiopod genus Lingula. Again it reveals a rapid rise in the number of species at the beginning, from the Cam- brian to the Ordovician, of the order of 40-50 million years. But between the increase and the decline phases, a more or less “‘station- ary’’ phase is intercalated, during which the number of species did not vary greatly. The decline phase of Lingula set in with the Car- boniferous. It is a long-drawn-out phase and continues into the present. Another interesting case is that of the coelacanths, a group of the fringe-finned fishes or Crossopterygii (fig. 4). This group never had a truly explosive phase. Nevertheless, there is an increase phase from the end of the Devonian to the Triassic, lasting about 100 million years. One more example may be given, the molluscan genus Poiretia (fig. 5). Here, the explosive phase in the evolution of the genus is 80-40 million years. Other examples confirm that the length of the explosive phase is of the order of a few 10-million years, the extremes so far found being 30 and 100 million years and the mean around 50 million years. As an example from the plant kingdom, illustrating at the same time a group with stable, geometrical characters living in an environ- ment which changes but little, diatoms may be shown. Small in- vestigated the chronology of their evolution, and the diagram of Hemiaulus shown (fig. 6) is a translation into our method of plotting of one of his diagrams. It shows a steepening rise to a climax in the Cretaceous. The increase phase lasted about 80 million years, no longer than in groups living in less stable environments, but there was a pronounced initial lag phase. Now it is interesting to consider some higher systematic categories. One might expect that the explosive phase in the evolution of higher systematic units, like families, orders, classes, is longer than in genera. But this is not so. 254 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 J eke) See He 50 fe) million years CRETACEOUS PALAEOGENE NEOGENE FIGURE 2,—Diagram of the numbers of species in the sea-urchin genus Salenia from the Cretaceous to the present day. (Surface area proportional to the number of species.) TATA 200 Tele) i 400 million years U.CAMB.ORD. SILUR.DEV. CARB. PERM, TRIAS.JURAS.CRET. CAINOZOIC Ficure 3.—Number of species in the brachipod genus Lingula, from the Upper Cambrian to the present day. (Surface area proportional to the number of species.) TIME IN EVOLUTION—ZEUNER 255 COELACANTHIDAE SRT RES TE 250 200 150 100 50 1@} Diplocercididae _ 300 million years DEV, CARBON. PERM. TRIAS. JURASS.CRET. CAINOZOIC Ficure 4.—Diagram showing frequency of species of coelacanth fishes in relation to time. Inthe Upper Devonian the family Diplocercididae flourished. In the Carboniferous it was replaced by the Coelacanthidae, which reached their climax in the Triassic. The small number of coelacanths recorded from the Permian may not represent their true frequency owing to the fact that relatively few marine deposits are known from the Permian. Possibly, therefore, the rise to the climax in the Triassic was not interrupted in the Permian, as is shown in the diagram, but was continuous from the Lower Carboniferous; if so, the rise would have taken about 100 million years and the rate of evolution would then be one of the slowest known. The coelacanths are regarded as a very con- servative group which changed but little in the course of time. 60 50 40 30 86.20 lO oO million years PALAEOGENE NEOGENE. Figure 5.—Diagram showing the number of new species of the molluscan genus Poiretia appearing in the different subdivisions of the Tertiary. The maximum number of new species was produced in the late Palaegene, within 30 to 40 million years of the appearance of the genus. 256 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The first example of this kind is that of the rotaliid Foraminifera (fig. 7), showing the number of existing genera in the superfamily. Again there is a phase of steepening increase. It lasted from the early Cretaceous into the Oligocene, for about 70 million years. Another case is that of the genera in the brachipod family, Spiriferi- dae, with an explosive phase of about 80 million years. A third example shows the orders in the class of the true fishes. This diagram 0 HEMIAULUS (DIATOMACEA) TOTAL OF SPECIES LIVINGIN ANY PERIOD NO. OF NEW SPECIES PRODYCEDIN ANY PERIOD Triassic ———> |Jurassic—o|L.Cret. |UpCretac|Eocene OL |Miocene|Prioc. | R. 200 MILLION [100 MILLION YEARS Figure 6.—The diatom genus Hemiaulus. The number of species living in any period (blocks) and the number of new species appearing in any period (curve) from the Permian to the present day. (Numerical material from Small, 1948. Ordinate scale linear.) 150 100 © 50 fo) million.years TRIASSIC JURASSIC CRETACEOUS PALAEOGENE NEOGENE Figure 7.—Diagram showing the number of genera in the superfamily Rotalioidea (Foraminifera) present in various geological epochs. (Surface area of blocks proportional to the number of species.) (fig. 8), which is based on Romer’s work, is not strictly to scale, but there is no doubt that orders like the sturgeons and gar pikes had explosive phases of some 20-60 million years. Finally, the number of orders within the class of the winged insects (Pterygota) must be mentioned. Since about a hundred orders have been distinguished, it is possible to plot them in much the same way as species in a genus. Once more, an explosive phase emerges, an exceptionally large number of orders appearing during the Upper Carboniferous and the Permian, over a period of about 60 million years. TIME IN EVOLUTION—ZEUNER 257 TABLE 2.—Insect orders Existing New 1 DYERICO LOVE OLE es as Toes Aes popes SEG we gs yee Adis Mallee erent yen aes 0 0 ower! Carboniferous ee sac te ete ae en eit a eee Pe 2 2 Wpper Carboniferous! =. a2. es 2S 25s oe nee cee aes ee 18 16 Perm is nes is eee eae eee Pe ne ees Bg 37 30 IMlESOZO1C =e Sree 2 ee Mee at gs tei iE E Ey Ti ib oem pe Ros Ye pe eS eee ep 31 22 CAS oy RN Mey een al RE te ape ree OnE ee et eet oe ey ge 38 13 BB eY 1s} 0) ee ey Se As ISR eS Vege ae aR eee EN Pore Se A PE aero SE CI Sea ee eC million ears | | aoe > : CAINOZOIC i i sul CRETACE- i y OUS $ 2 = eo 7 NG JURASSIC 3 of °. 8 : on | jee 4 PERMIAN B, : 5 CARBON : : ‘ -IFEROUS DEVONIAN SILURIAN FiacurE 8.—The evolution of the true fishes correlated with the geological time scale. Based on a diagram by A. S. Romer. To summarize, increase phases (often of the ‘‘explosive” type) in the evolution of genera, families, and orders lasted, so far as evidence goes, something like 15-100 million years, with a mean of about 50 or 60 million years. In other words, the increase phase of evolution is of the same length [of the order of 50-60 million years (+30)], irrespective of the sys- tematic unit investigated. It is difficult to resist the temptation to speculate on the significance of this result. What it appears to mean is that the number of species steps involved in the evolution of new systematic categories is the same irrespective of the rank of the cate- gory. It takes no longer for a new order to evolve than for a new genus to appear. 258 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 If this is accepted as a valid inference from the material studied, then the difference between genera, families, orders, and classes must lie in the quality of the unit steps involved in their evolution. This is not a new idea,? though we have arrived at it from a new and un- expected direction. What is meant by the statement that the quality of the steps de- cides the systematic category which is evolving, is easy to illustrate but difficult to formulate. A good example is the evolution of the jaws of the fishes from gill arches. This is an extraordinarily interest- ing case of change of function of an organ. Its real significance, however, is that this change provided the fishes with a decided ad- vantage over their environment. It opened up new and better food supplies and, as Sewertzoff expresses it, “increased the general life- intensity of these animals.”’ A biting-mouth skeleton is important also as an offensive and defensive weapon. It is, I think, sufficiently evident that this particular change of function was full of evolutionary potentialities. Applying Sewertzoff’s terminology, such changes may be called aromorphs. On the other hand, if one considers the extreme adaptation of a leaf insect (Phyllium), one finds that it has—to use Sewertzoff’s phrase- ology—led to a decrease in the life-intensity of the animal. No new food supplies have been made available by the process of adaptation, and its evolutionary potentialities are nil. This type of adaptation is typical of the lower systematic categories. There is another interesting feature that. emerges from the chrono- logical treatment of evolution. It is that there appears to be no correlation between fast rates of species evolution with groups having a rapid succession of generations. Elephants are among the most slowly breeding animals known and yet their rate of evolution was the fastest so far found. On the other hand, Drosophila, which, because of its rapid succession of generations, is so much used in experimental biology, is a genus 50 million years old. It appears therefore that a rapid succession of generations must not be taken as a, substitute for long periods of time. It is indeed surprising to find that the number of generations is not the only factor ruling the rate of change in evolution and that this change is in a vague way correlated with absolute time. Does this perhaps mean that external, environ- mental factors are influencing evolution over very long periods of time? I am unable to answer this question, or to offer any other explanation. I hope I have been able to show that a study of the chronology of evolution is well worth the effort. The suggestions made here must ?This has been put forward in different forms, for instance, by G. G. Simpson and R. Goldschmidt. TIME IN EVOLUTION—ZEUNER 259 not be taken as final results. For the time being the material is still too scanty to make generalizations sufficiently safe. Some of the time rates, however, have been found by several workers inde- pendently, and some of the rules—if one be allowed to use this term— have been deduced by more than one worker. BIBLIOGRAPHY GoLpscHMIpT, R. 1940. The material basis of evolution. 436 pp. New Haven. ScHINDEWOLF, O. H. 1947. Fragen der Abstammungslehre. Aufs. Reden Senckenb. Nat. Ges., Frankfurt a M., vol. 1. 23 pp. Simpson, G. G. 1944. Tempo and mode in evolution. 237 pp. New York. SMALL, J. 1946. Quantitative evolution—VIII. Proc. Roy. Irish Acad., ser. B, vol. 51, No. 4, pp. 53-80. 1948a. Quantitative evolution—IX—XIII. Proc. Roy. Irish Acad., ser. B, vol. 51, Nos. 17-21, pp. 261-346. 1948b. Some laws of organic evolution. 15 pp. Privately printed. ZHUNER, F. E. 1946a. Biologicai evolution and time. Ch. XII in Dating the Past. London. 1946b. Time and the biologist. Discovery, vol. 7, No. 8, pp. 242-249, 256. (Some diagrams shown in the discourse are figured in this article, and acknowledgments are due to the editor of Discovery for per- mitting their reproduction.) he sabi Bry) “SS ay -_ iL ° fo — : a han Fhe ae BS Aydihiet pond Win! : idee organ 4 anibe tne ENE “spe pie rie rh eta) iD ; pone i 9n0 rn anon Noe cgens vd 4 wl # Th ’ ; ; s i ee : ie oe a +3 “=F crits Myf 17 f i Vee \ ¥ _ ‘ ii srry ‘9 ve a ‘ 3 mr - = _ , pormliwat wih) .cubiwloys Wn siead Wsquieit alt O45 Z re : : , 5 Z| : ‘ ‘ : +t us s Kf% era a eat) pan deine ce ack ailotegati iets Sal eg TE : A . ty On eh HO? 995 a mbt —— hs _ S t a ; B 7 i ee “= Pada) CRA a” ; ‘ | ¥ yas any BY wks btee ds Bh. oheadiz ii er "i — fs | | ; . ppt ee: sy A A600 Dah botal. tL, anier 1 e74 f—aohuior ave ii avid ae a : eae he " e-88 AS * an 0 —_— L are ie el fa A heel’ ; (Or er TELS A edt if "5 by aaalstta nity? dope eh ibe 5: iar’ OG ae vega TS Df dial Soe < ; Sue. « Shade 3 op iat 1) f f fio i Add CASAS . rel adres oe be ir. ine : ' - il ni : ‘. : ; : ca A tons : , EAS. hee a ps hs - r tk , oe rive: Menten TLE Le Lee ir } with) hia neil toys uotyclotl. evel | a : ag a Sf « ip a a o,f \ Par) he i Ni sta, At : Ls rte 4 | : one al 7 ‘a hl Riri | bad bia at Us ie ania ‘edity pill te Sig a beats rw riyais Sine ne ‘ets sai ae ih TOR see sada Cr Ye ow Vw ii wel ; writs! tte i gaks a) = r j i» (ape ¥ ee 43) bia i sada! 12 399% MORE ABOUT ANIMAL BEHAVIOR ! By Ernest P. WALKER Assistant Director, National Zoological Park [With 16 plates] INTRODUCTION The scope of the subject ‘‘animal behavior’ is almost unlimited, for every species has its own particular pattern of behavior and the indi- viduals of the species show many variations even from that pattern, so that the subject could really be made an exhaustive study for each and every species. Although psychologists have made extensive studies of human behavior, actually they have merely made a good beginning. Glimpses of animal behavior and some conception of how behavior patterns may have developed frequently help in explaining human behavior and human reactions, for our reactions are the result of a long series of experiences of trial and error and elimination by natural causes just as has been the case with those of other animals. Similar causes have brought about similar reactions in many instances, and different causes and environment have developed different reactions; there is, therefore, a remarkable diversity of behavior pattern among the different kinds of animals that live under many different conditions. Those that have not been able to adjust themselves to their environment have become extinct and are usually known only by their fossil remains; those that could adapt themselves and multiply are the forms that have survived to the present day. The activities in an animal’s life are essentially the same as the basic activities of humans. Individuals must survive, and to do so they must be able to avoid enemies, to obtain food and shelter; and for the species to survive, the individuals must reproduce and the young must be given an opportunity to start their own struggles for survival. No doubt everyone who has observed animals closely has seen them do many things that could not be explained in the light of our ex- periences and practices. However, we must judge and interpret the 1 An earlier article by the same author, under the title ‘‘Animal] Behavior,’ appeared in the Annual Report of the Smithsonian Institution for 1940, pp. 271-312, 18 pls. As the first paper has been out of print for several years, some of the same ideas and examples contained in it are incorporated in the present paper, although not as exact quotations. 261 262 § ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 actions of animals in the light of the lives they normally lead in the wild. If we are well enough informed as to the conditions under which they live and their mode of life, we can often understand actions that would otherwise be incomprehensible or aimless to us. I do not believe that animals do any more aimless things than humans. On the contrary, I am often impressed by the direct and efficient manner in which they carry on the work of their lives. The habits of animals are so intimately associated with and gov- erned by their structure that one who is somewhat familiar with either the habits or the structure can often make rather close deductions from one regarding the other. Every habit or type of behavior that has been developed because it helps the animal in its life is, of course, a special trait. Some are so little known that attention might be focused upon them and con- sideration given to the manner in which they benefit the animal. To understand the animals best, we must realize that merely because we cannot perform a given act does not mean that an animal cannot do it. For example, the sense of smell in most humans is so unde- veloped that we cannot trail another animal by scent, as we know many animals do. Also, our hearing is so dull we cannot detect sounds that we know many other animals hear. Likewise, other senses of animals are probably so much more keenly developed than ours that they re- celve impressions or information of which we know little or nothing. For example, some animals give off vibrations that other animals are able to detect; the presence of a warm live body can be detected by certain snakes at distances of several feet even though their eyes and nostrils are not functioning; and apparently impressions are received by insects through their antennae and by fish through their lateral lines. How individual animals acquire their behavior patterns is a fas- cinating field for study. Some actions are apparently taught to the young by the parents; some are learned by the young by observing others of their kind; some are learned by the trial and error system, or as we know it, “by bitter experience”’; and some come to the animal by instinct; that is, the animal reacts in a certain way (usually the right way) under certain circumstances without previously having had an opportunity to learn consciously to act in that manner under those circumstances. How such reactions develop is explained by various theories and is a separate study. ECOLOGY, STRUCTURE, AND HABITS Ecologists refer to an animal as occupying an ecological niche, that is, the main activities of the animal take place within a certain type of habitat. It may live entirely within the water, on or under the ground, or almost entirely on trees, or combinations of any of these, ANIMAL BEHAVIOR—ERNEST P. WALKER ”263 and its food is definitely limited by the products that it can obtain in the habitat which it occupies. Animals that do not use the same food or are not antagonistic, because one does not prey upon the other, can occupy the same areas or overlapping areas, so that ecological niches do not have definite, clearly defined boundaries, but are rather areas or space spheres of activity having also a time component. The structure of the animal to a large extent governs the type of ecological niche that it may occupy, since through evolution the animal’s form has been modified to adapt it to the type of ecological niche that has been available to its long line of ancestors down to itself. Thus we can to some extent explain and understand the great variety of forms of animal life. We might consider two little creatures of approximately the same size which occupy different ecological niches and are very different in structure, and see how they fit into their respective spheres. These are golden hamsters (Mesocricetus auratus) and “‘flying’’ squirrels (Glaucomys volans). The golden hamster of Syria averages about 4 ounces in weight, with a maximum of about 8 ounces. It is a stout-bodied, short-necked, short-armed, and short-legged little mammal with short fingers and toes and a tail only about half an inch in length. It has very large cheek pouches that open inside the lips and extend far back of the shoulders, in which it can carry surprising loads of food or nest mate- rial. It is rich golden-reddish brown above and white beneath, with white hands and feet. The little creature is an inhabitant of an arid, rocky region where it lives mainly among rocks or in burrows in the ground around the rocks. Syria’s climatic conditions result in a scarcity of food material for such little creatures for long periods of time; therefore, the storage of food is necessary and the cheek pouches are of great value in carrying food to its burrow or den. It works industriously at this, apparently the only limit to the amount of food it carries being determined by the amount that is available. They are relatively slow-moving, clumsy little creatures in com- parison with many other animals that we know, but are obviously well adapted to leading their lives in an efficient manner in their habitat. Their movements remind one of military tanks in that they are slow, ponderous, and persistent. There is nothing sprightly or agile about them, but if they want to go in a certain direction or up a certain surface they keep at it so persistently that frequently they succeed, and they can wedge themselves through surprisingly small crevices. They display remarkable persistence and determination to get into crevices that would seem to be so small and uncomfortable as not to be attractive to them. The mouths of rodents are definitely on the underpart of the head and back of the nose, making it difficult for them to cut upward when 866591—50-—18 264 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 in their normal position. The animals solve this problem very readily, however, by throwing themselves on the back or side and cutting upward with their teeth. A picture of such an effort is shown in which my “Shehammy” has thrown herself on her back and is trying to cut upward on the lower edge of a door so that she can squeeze through the crack between it and the floor (pl. 1, fig. 1). This aperture was only seven-eighths of an inch high and, when she tried to force her way through it standing on her feet, she was much too thick and her body was much too tense, but in the inverted posi- tion she extended her arms forward so that she was able to get a grasp on the lower edge of the door and pull herself through the crevice, which she did repeatedly. This ‘““chammy,”’ one of my pets, did it regularly, and I have seen other rodents do it occasionally. Frequently one of my golden hamsters, when permitted to run about my study in the evening, would clamber up in the crack between a bookcase and the wall a foot or more above the floor and stay there for a considerable time. She usually rested by bracing her feet against the wall and her back against the bookcase and apparently was content to stay in that position. This suggests to me that these animals adopt such a position as a means of being comfortable in fissures between the rocks among which they live. When hamsters want to get down from an elevation that they have reached, they apparently do not think of jumping down even a few inches, but lower themselves as far as possible with the hind feet, sustaining themselves by the top of the hind feet, which are bent at the ankle. Next they release one foot and then the other and fall head downward, usually landing on their noses, but sometimes on their backs (pl. 2). They then sit up, sneeze, rub their noses, and go about their business. It will be noted that they do not utilize their hind claws or toes which are too small to be effective. This method, while probably a relatively inefficient action, has persisted because it is not seriously harmful in the circumstances under which they live, and is the best they can do with their short fingers and toes which are not of much use to them for such descents. At night, when foraging for food, hamsters apparently spend very little time eating, preferring to fill their pouches, take the load of food home, and return for more (pl. 1, fig. 2). In this way they gather food when it is available and eat at leisure in their nests without being exposed to danger. During the daytime, their sleeping period, they frequently wake up and eat, probably consuming much more food at this time than during the night when they are actively foraging, grooming, exploring, or visiting their neighbors. When “‘hammies” are away from their homes or other shelter and there is a sudden alarm, they dash for shelter with surprising speed for little folk with short legs but if they are touched or attacked before ANIMAL BEHAVIOR—ERNEST P. WALKER 265 they can get away, they throw themselves on their backs incredibly quickly and are prepared to fight savagely with their sharp, fairly strong teeth and tiny claws. If they are in their nests, they pay little attention to outside disturbances, at most sniffing from just inside the doorway. Baby ‘‘hammies” arrive in litters of as many as 17, tiny, blind, naked, pink, helpless little fellows that lie on their backs and nurse while mother “‘hammy” hovers over them to keep them warm in the snug cup-shaped nest of fibers she has constructed in a remote portion of the burrow or den. The little ones grow very rapidly, and by the thirteenth day they have a good coat of fur, are eating solid food, and begin wandering about in the burrow, den, or cage. Even though their eyes are not yet open, they find their way back to the nest if mother has not detected their absence and carried them home. By the fifteenth day their eyes are open, and by the seventy-third day they produce babies of their own. ‘They bear large families in rapid suc- cession but cease to breed regularly when about 1 year old, and the life span is short (probably a maximum of about 3 years). There is a high mortality rate, probably throughout life. The structure and lives of the flying squirrels (Glaucomys volans) of North America are in marked contrast to those of the hamster. Flying squirrels weigh about 3 ounces, have slender bodies, long, slender, strong arms, fingers, legs, and toes with fairly long, curved claws, and a rather long tail. Theskin of the body is much larger than is necessary to enclose the delicate little form and is extended outward along the side as hair-covered membrane indistinguishable from the skin of the body. This skin reaches down to the wrists and ankles, so their outline is almost square when the little creatures put their arms forward and out and their legs back and out as they do while gliding through the air. This adaptation more than doubles the area of the upper and lower surfaces as it appears when the animals are at rest. In keeping with this form, the hairs of the sides of the tail are long, and closely set, and project at almost right angles, whereas those of the upper and lower surfaces are very short and lie close to the bone of the tail, so that the tail is, in cross section, almost like a feather. All these are modifications for life in an entirely different habitat from that of the hamster. Flying squirrels live among the trees, generally making their nests in hollow limbs, old woodpecker holes, or other shelters high above the ground. They are strictly nocturnal, having very large eyes adapted to admitting the maximum light available. They are not limited in their movements to climbing up and down the trees or to mere leaping between objects, for their remarkable form gives them power to glide long distances. ‘To do this they seek an elevated point, leap out into space and spread their arms and legs so that they take 266 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 on the form of miniature gliders (pl. 4, fig. 1). When approaching a landing, which they generally try to make on a tree or other object not on the ground, they swing slightly upward to check the glide and bring their long arms and legs forward as shock absorbers (pl. 4, fig. 2). Instantly upon alighting they dart around to the other side of the tree or limb, which is probably a provision to escape any enemy that might be following them. Before flying squirrels take off on a glide they almost invariably sway the head and body as far as possible first to one side then the other, repeated several times, and often raise as high as they can stand and crouch as low as possible. 'Thisis probably a sort of range finding by triangulation. They feed very largely on nuts and acorns which they gather and store in their nests and probably in almost any location in the forest in which they can find a cavity that is large enough to hold a nut securely. I assume this from the fact that my pets regularly place nuts at many places about the house other than in their nest box. They particularly selected the top of the window, the medicine cabinet in the bathroom, my hand, my pockets, inside my collar, and even the depression between my arm and body when I have my arm close to my body. When placing nuts, they have an interesting habit of trying to make them secure wherever they are leaving them. They force them into the crevice or onto the surface where they are leaving them, then tap them three or four times with their front teeth. This suggests that they probably similarly try to wedge the nuts in cracks in the bark or slight depressions on the tops of tree limbs in the wild. In addition to nuts and seeds, they eat some fruits, berries, and insects. My pets eat mealworms, grasshoppers, and small bits of meat in moderation. “Glauckies” (short for their scientific name) are very hesitant about going down to the ground, feeling much more at home leaping about in the trees, gliding from place to place; however, they will go down on occasion. When they do so, they are plainly not at home and run with the arms and legs extended so as to hold the body as high above the ground as possible, the gliding membrane being pulled upward close to the body so that a very peculiar effect is produced. Instead of sleeping soundly and ignoring disturbances in their neighborhood, as do the hammies and many other burrowing creatures who know they are safest in their dens, the flying squirrels sleep lightly and at the least disturbance in their neighborhood they peer out or dash away. “Glauckies’”’ are born in families of two to six pink, helpless little ones, with the gliding membranes plainly evident. They develop ANIMAL BEHAVIOR—ERNEST P. WALKER 267 slowly, are not well clothed in fur until about the twenty-eighth day, and continue nursing until about the fifty-sixth day. In recognition of the dangers of an active life in trees, they, like young tree squirrels (Sciurus), start out exploring near the nest with great caution, and venture farther and take more chances only as they gain strength, experience, and confidence. Their gestation period is about 60 days and young are born when the parents are about 1 year old. These brief descriptions of the widely different hamsters and flying squirrels, which inhabit very different ecological niches, give a glimpse of the specialization among animals which enables them to survive and to lead their lives under diverse conditions. Giraffes (Giraffa) have developed such a high degree of specializa- tion that they are able to fill an ecological niche without competition with other mammals except in those portions of their range which overlap that of the elephants. Much of the food of the giraffe is taken from the upper portions of shrubs or from the lower branches of trees far above the reach of most mammals. In addition to its long legs and very long neck, the giraffe has elongate, mobile lips and can extend its tongue several inches to help it gather in leaves and twigs. The moles (Scalopus and other genera), which burrow in the earth, are blind, have exceedingly short forelimbs, with nose and tail very sensitive to touch, large, powerful hands armed with long, strong claws, and very strong muscles in the fore part of the body, particularly those associated with the arms. The fur is very soft and short and will lie in any direction; thus it does not impede the mole when it desires to run backward in its burrow, which it often does. Moles feed on earthworms, grubs, and other small animal life they catch in the earth. (See pl. 5, fig. 2.) In marked contrast are the gibbons (Hylobates and Symphalangus), highly specialized members of the primate or monkey group. Their arms are exceedingly long and powerful, with elongate hands and four strong fingers, the thumb being small and situated far back on the hand. The legs are rather long, and the feet are better fitted for grasping than are the hands. The body is small. These adaptations enable gibbons to be the most expert of all mammals in swinging by their arms through the forest from branch to branch and even from tree to tree. In this process the hands are used as hooks, the thumb taking no part in grasping. Since the hands are usually occupied when traveling through the trees, gibbons carry objects such as food or branches by grasping them with the feet. The same method of carrying objects is used by chimpanzees (Pan) and orangutangs (Pongo). Whereas the gibbons, chimpanzees, gorillas (Gorilla), and orang- utangs lack tails, the spider monkeys (Ateles) of Central and South 268 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 America possess long, prehensile tails which they use in many ways, especially to hang by or to use as a supplementary hand in steadying themselves. In the picture (pl. 11) it will be seen that one of the animals is using its tail to hold itself while it leans far forward. On one occasion I saw seven spider monkeys in this group leaning forward, held by their tails, watching for the door to open so they could go into the house for their afternoon meal. Some rodents have large, powerful incisor teeth that protrude much more than those of the majority of rodents, and one naturally surmises that this trait is associated with some habit peculiar to them, which indeed it is. The animals possessing such teeth use them in burrowing; that is, they cut away the earth, remove small stones, and cut roots, thus performing most of the functions that the powerful hands of the mole perform and, in addition, do the cutting work which the mole’s hands cannot do. Some other rodents, and many of the insectivores also, have projecting incisor teeth, but these are relatively small and slender and are obiously not suitable for such heavy work as digging, moving small stones, or cutting. These are used as forceps for the picking up of food, which consists mainly of small fish, insects, or worms. Such animals as the kangaroos (Macropus), the kangaroo rats (Dipodomys), the African spring hare or spring hass (Pedetes), the jerboas (Alactaga, Scarturus, Salpingotus) and others have large, long, powerful hind legs, relatively small, weak, short forelegs, and long tails which are either tufted with long hairs near the tip or are rather heavy, often very thick near the base (pl. 5, fig. 1). These animals are leapers and use the hind legs almost exclusively in propul- sion. The tail is a balancing member; it also acts as a tripod leg in some forms. The types of leaps of different animals vary greatly. One interest- ing difference is the contrast between the leap of the flying squirrel, an animal that leaps directly toward its objective with a very flat trajectory, often planning to land below its target and then climb up to it, and the leap of the tarsier (Tarsisus) which has a very high trajectory and lands the leaper at its objective. After becoming accustomed to photographing the leaps of flying squirrels, I tried photographing tarsiers leaping and aimed my camera as I would for the squirrels. With the pronounced leap upward of the tarsier, I at first caught only the tail and hind portion of the animal in the upper portion of the picture. After a few such experiences I learned the difference in their leaps and corrected my aim accordingly. The list of such highly specialized forms could be expanded indefi- nitely to include representatives of all groups of animal life, for every animal is specialized to lead a certain type of life. ANIMAL BEHAVIOR—ERNEST P. WALKER 269 CLEANLINESS Almost all animals are fastidiously cleanly about their persons and their homes. Because of the fact that many animals live on or in the ground or on trees or other surfaces that frequently are dirty according to our standards, we often erroneously consider the animals to be uncleanly. As a matter of fact, practically every animal takes great pains to avoid soiling its coat, and when it does unavoidably become soiled, the animal at the first opportunity painstakingly cleans itself by shaking, rolling in the sand, washing in water, or licking itself. Even those pests, the common Norway rat, roof rat, and house mouse, which have found that they can obtain food and shelter in man’s filthy surroundings, endeavor to keep themselves clean. If they are observed when not alarmed, it will be seen that they carefully pick their way through their surroundings in an effort to avoid soiling themselves, and after they have finished their exploration they will invariably be seen to groom carefully. If one will part the fur of small mammals it will be seen that the skin is generally immaculately clean. A star-nosed mole (Condylura cristata) that I am keeping as a pet comes out of the ground, goes into a dish of water and at once begins grooming. In a few minutes it has washed thoroughly. Animals have developed many ways for keeping themselves and their nests clean. Apparently flying squirrels rarely void excreta during their entire sleeping period, which may be as much as 16 hours. Obviously they wish to keep the inside of the tree or other nest loca- tion clean, so wait until they go out at night when they can see clearly and there is a minimum of danger from outside enemies. Many of the burrowing mammals have little toilet rooms that open off of the main burrow where they deposit their excrement, thus keep- ing the main burrow clean. Young North American opossums (Didelphis virginiana) of an age to be in the pouch or still be clinging to their mother apparently do not release their excretion unless they are stroked on the lower abdominal region. This is probably a pro- vision for keeping the pouch clean until such time as the mother is ready to clean the little ones. POSTURES The new-born babies of many of the small mammals naturally lie on their backs. This leaves them in the proper position for nursing when the mother hovers over them, protecting them, hiding them, and keeping them warm. Most hoofed animals do not voluntarily sit on their haunches. A horse assumes this position briefly when getting up, but this does not constitute true sitting. The exceptions to the rule are the tapirs, 270 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 which frequently sit on their haunches with the forefeet supporting the front part of the body (pl. 14, fig. 2). A position that would appear to us to be uncomfortable is regularly adopted by flying squirrels, which normally live in hollow trees. The body hangs in a vertical position, either head up or head down, sus- pended by the fingers or the toes, most of the load actually being on the fingernails or the toenails (pl. 3, fig. 1). They sleep in such poses at least a portion of the time. The hanging position is probably assumed as the result of having accommodated themselves to situations where there is no satisfactory ledge or bottom in the den, so that a tree cavity that would otherwise be unsuitable can be utilized by clinging to its rough surfaces. However, they now take this pose freely even when comfortable nests or rests are available. Of course, they also take other positions such as curled up in a ball or lying on the side or the back. The inverted position so often used by sloths (Choloepus, Bradypus, and Scaeophus), wherein they hang beneath a limb, is well known and is entirely suitable for their purposes. Bats generally hang head downward. HOMING INSTINCT When an animal has become established at a given location, it almost invariably develops a strong attachment for its home even though such home does not provide entirely satisfactory quarters. Almost every animal has a rather definite area in which it lives its entire life, and it rarely leaves this range except when forced to do so by very unusual circumstances. This is commonly shown when captive animals escape, for they usually remain nearby for some time and frequently go back into the cages if they are able todo so. People who are familiar with animals regularly take advantage of this trait by leaving the cage door open, refraining from frightening the animal, and placing food in the cage or nearby until the animal is recaptured. On one occasion I was keeping a specimen of the least short-tailed shrew (Cryptotis parva) in a jardiniere. I had observed it trying to leap out, but it is not adapted to leaping and was apparently unable to reach the top. After a few days I discovered that it had escaped during the night probably by superior jumping gained by persistent exercise. It was so tiny that it was useless to search for it in the house, so I waited until evening and then placed the jardiniere on the floor almost directly below the place at which it had been standing on my desk. JI crumpled around it an old blanket so that it formed a ramp or runway from the floor up to the top of the jardiniere. I then mashed a mealworm and dragged particles of it in a trail on the blanket from where it touched the floor to the top of the jardiniere. Within a couple of hours the little fellow was back in his home. ANIMAL BEHAVIOR—ERNEST P. WALKER 201 Plainly he had come to recognize the jardiniere as home and was glad to get back to it when it was made possible for him to do so. Remarkable developments of the trait of returning to the home range are shown by carrier pigeons (Columba), and birds, mammals, and fish that migrate. SELF-PRESERVATION Humans ordinarily give little thought to the matter of self-preserva- tion, although it becomes a vital subject during wartime. Among other animals it is ever foremost. In its broadest aspects self-preserva- tion involves not only active and passive resistance to enemies, but also the ability to obtain food and shelter. Humans have recently had brought to their attention the fact that they, like other animals, must either be prepared to defend themselves or run away when an ageressor attacks or threatens. If there is no place to which they can escape, or if they cannot successfully fight off their enemy, they have only the alternatives of being subjugated and made slaves, as some- times happens among humans, or of being destroyed, which in the case of animals may mean being devoured. There are very few animals that are aggressive to the extent of trying to drive away or subjugate their neighbors merely for the pleasure of the victory. Ordinarily, they will start a fight only when it is necessary to obtain food, shelter, or a mate. The methods used by animals to avoid, escape, or defeat an enemy are almost as numerous as the species of animals, for the individuals of every species employ one or more methods. All are interesting and some are very remarkable. The seeking of shelter is the most common of passive defensive measures and is practiced to a greater or less degree by all animals. Some go into burrows in the ground, crevices in rocks, or holes in trees, while some of the larger ones retreat into dense jungles or forests. “Freezing,” the act of remaining motionless, is extensively employed, as at the first intimation of danger almost every animal ceases prac- tically all movement. Since an object which is very conspicuous when moving is readily overlooked when stationary, “‘freezing’’ is a highly successful means of avoiding detection by enemies. Even those that depend mainly on their ability to scent their prey may have difficulty locating it if they cannot see it. Rabbits and hares (Leporidae), quail (Colinus), and grouse (Tetra- onidae), when perfectly still in their native haunts, are very difficult to see, and often will not move unless one almost steps on them. This method of escape is applicable in almost any type of surrounding, even on seemingly barren plains or expanses of snow. It is most effective, of course, if the color pattern of the animal is such that it 272 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 harmonizes and blends with the surroundings or if the outline of the animal is obliterated by some type of camouflage. Most animals are so colored that they blend perfectly with their surroundings, some, such as the ptarmigan (Zagopus), certain hares (Lepus), and weasels or ermine (Mustela), even changing to white in winter to blend with the snow. Among insects, camouflage and mimicry have been developed to a high degree of perfection in many species. Some resemble their natural surroundings so closely that they are difficult to distingusih even though one is looking directly at them. Resemblance to dead or green leaves or stems is the most common camouflage, the pattern and form of the insect being modified to a strikingly perfect imitation of the color and form of those parts of the plant on which the insect usually lives. In the type of specialization known as mimicry, one species of insect very closely resembles and usually behaves like another which is distasteful or dangerous to animals. By this resemblance the imitator is often let alone by an anima) that mistakes it for the undesirable insect which it resembles. When it is not possible to avoid an enemy, some animals use @ more or less passive method which is well illustrated by the armadillos (Dasypus, Tolypeutes, and related genera) which roll themselves into a ball, completely protecting the feet, tail, and head within the armor plate of the body (pl. 6, figs. 1 and 2). This method effectively baffles many animals that would ordinarily devour them, and when the would-be attacker becomes discouraged and leaves, the armadillo unrolls itself and goes about its business. Another method is the feigning of death, which is well illustrated by the American opossum which, when alarmed, falls on its side with the mouth partially opened and appears so limp and inert that it is often left for dead by animals that would vigorously attack if they surmised the animal to be alive. Apparently opossums are unpalatable to many animals, so that this means of protection is very effective. When danger ceases to threaten, the opossum gradually resumes activity, but if the attacker is merely waiting nearby and makes a movement, the opossum will usually again go into its death-feigning act. It is supposed that this behavior is a type of fainting induced by fright and is perhaps not actually under the control of the animal. When the hog-nosed snake of the eastern United States, frequently called spreading adder (Heterodon), feels itself in danger, it feigns death, throws itself on its back, and assumes various grotesque poses if it is seriously aggravated. It does not, however, do this unless the annoyance is great and continued. Many insects feign death when a disturbance occurs in their vicinity. This is illustrated by the many beetles that inhabit low shrubbery, which drop from their elevated locations to the ground and remain quiet or quickly take refuge in a more protected location. ANIMAL BEHAVIOR—ERNEST P. WALKER 273 One of the most remarkable modifications of feigning death of which I have heard was witnessed and described to me as follows by John N. Hamlet, of the United States Fish and Wildlife Service: Three of us recently saw a Cooper’s hawk (Accipiter cooperi) chase a spotted sand- piper (Actitis macularia). The piper dropped into the water and stayed under for several seconds. The Cooper’s lit on the stream edge a few yards down the stream. The sandpiper came to the surface and floated down the stream with its wings open flat on the water and its neck stretched out. It passed within 3 feet of the hawk who gave it no more than a casual glance. The piper floated down stream about 20 yards and took off and disappeared. The live sandpiper drifted close by the hawk but was not recognized because it was not moving in its usual pose. This is a choice example of the effectiveness of ‘freezing’? and assuming an unusual pose. Mr. Hamlet has witnessed similar action by sandpipers on two other occasions. Bluff is another effective defense that is employed by many animals. Its best form is for the animal to face its enemy in a pose not usually assumed by it and that makes it appear as large as possible, ferocious, and threatening. Many, if not all, of the owls (Strigiformes) bluff by crouching low, spreading their wings at almost right angles to the body and ruffling the body feathers until the bird appears several times larger than it really is. The bittern does likewise, and most mammals bluff to some degree. A good example is the dwarf weasel (Mustela rixosa campestris), only about the size of a cigar, who stands his ground, opens his mouth wide, barks, and even attacks if need be, although its teeth and jaws are so small it can scarcely break the skin of one’s hand. The domestic cat’s (Felis cattus) high-arched back, bushed-out tail, and wide-open, snarling mouth present a good example, and many of us have witnessed the hestitation of a dog suddenly confronted by such an attitude. Often this hesitation gives the cat an opportunity to escape without having to fight. Fighting for mates is definitely beneficial to the species, for by nature’s law of the survival of the fittest, which must prevail through all species, only those survive that are best able to take care of them- selves. This, taken in its broadest sense, includes not only physical strength but mental alertness and adaptability to varying and new conditions. Males that are physical weaklings and would not father vigorous offspring are ordinarily vanquished in encounters for mates and therefore leave no progeny. By this process nature has consist- ently eliminated the unfit and has improved each of the species. The occasional maiming of individuals in their conflicts is not in itself injurious to the species, as such mutilations are not inherited and, if the parent was vigorous, the progeny stand excellent chances of being vigorous even though the parent may have been injured in some of its conflicts. 274 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Among animals that live in colonies or in herds, conflicts are gen- erally rare except among males that are fighting to determine who shall be master of the herd, as in the case of bison. However, on one occasion I witnessed a definite fight in a prairie dog (Cynomys) colony. In this case the aggressor was outside of the burrow and trying to keep another one in the burrow and, at the same time, to fill the burrow with dirt to bury him. Several others were within a few feet watching the proceedings but taking no part in it although it was obvious that they sympathized with the “‘underdog.’”’ Whenever the one that was in the burrow would attempt to come out, the one that was outside would try to bite and scratch him, and, when he had him forced back into the burrow, would scratch and push earth into the entrance. Finally, after perhaps 20 minutes, he lost interest and left. The bystanders almost immediately went to the burrow, greeted and apparently sympathized with the one that had been attacked, and in a few minutes the normal life of the colony had been resumed. This method of fighting by trying to imprison the victim is also used by prairie dogs in closing the entrance to the burrow which a snake has entered. ADAPTABILITY The readiness with which animals accept approaches or friendliness of man varies greatly. Some seem to be so thoroughly imbued with caution and suspicion of man’s intentions that they can be tamed only with the greatest of effort. Others respond almost immediately to handling and friendly treatment. Examples of the latter are the gray foxes of the United States (Urocyon) which tame very readily and in marked contrast to the general wariness and slow taming of the red foxes (Vulpes). The beaver (Castor) is another animal that tames easily, sometimes merely a few hours of kind treatment being sufficient to win its confidence. Young hair seals (Phoca) also tame almost immediately when captured. Usually they seem to have no fear when picked up, looking to their captors as friends and becoming affectionate pets, sometimes swimming after a boat that is leaving them behind after its occupants have picked them up and petted them briefly. Penguins are fearless and very curious as to visitors on land or on the ice or snow where they normally have few or no enemies, but in the water they are cautious as there they are accustomed to watching for enemies such as killer whales (Orca), sea leopards (Hydrurga), and large fish. Another phase of the adaptability of animals is the degree to which wild animals can survive where man has established himself. Most of them appear to have no fear of man’s presence and his activities so long as they are not actually molested and their haunts and food ANIMAL BEHAVIOR—ERNEST P. WALKER 275 supply are not destroyed. We can see numerous examples of animals that have gone ahead fairly successfully with their ives when man has not interfered too much with conditions essential for them to live. Another group consists of animals that thrive in close proximity to man and either become a part of man’s household and receive his direct aid in their existence, or adapt themselves to the conditions that man provides and obtain food and shelter in spite of his utmost efforts to control them. In this group are the Old World rats (Rattus) and mice (Mus) that have become established almost throughout the world wherever man has made settlements. Other examples are the European house sparrow (Passer domesticus), commonly called English sparrow in the United States, and the European starling (Sturnus vulgaris) both of which have become firmly and widely established in North America. These two birds, like the rats and mice, have be- come pests in some regions where they have adapted themselves to a remarkable degree and have become very plentiful. They have found sufficient food and shelter around man’s activities so that they have thrived where other less adaptable forms have not been able to sur- vive. Indeed, in many cases they fight the native birds and take over the ecological niches normally occupied by the local birds that are less adaptable and aggressive and have not been successful in defending their territory. It is sometimes said that there is a third group of animals comprising those that cannot tolerate proximity to man but, if we study the problem, we usually find that man’s activities so changed conditions essential to them for food or shelter that they could not survive, or else that man intentionally killed them off. Most animals, if given to understand that man will not harm them, will become tame, and if man will feed them it is surprising how friendly even very wild kinds will become. I have a pet big brown bat (Eptesicus fuscus) that in a few days became so tame it would return to me after each of its flights about the room (pl. 13, fig. 2). Even when I awaken her she makes some effort to overcome her stupor and come into my hand to be warmed preliminary to taking her evening flights and receiving mealworms, of which she is very fond? Chipmunks (Zamias) that have been tame during the summer or fall, then go into partial or complete hibernation and are inactive for a period of from 5 to 6 months, come out in the springtime and resume their friendships and friendliness practically as though they have not been interrupted. One individual that came in the window onto my desk and was so tame that I took a picture of it while it was on my hand obtaining food, returned the following spring and re- 7 fa the Saturday Evening Post for February 4, 1950, is an account of further results of my studies of ats. 276 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 sumed our relationship where it had broken off with the beginning of bad weather the previous fall. Another example of the confidence of wild creatures is the case of two flying squirrels that are greatly loved guests in our home. They have such fondness for both my wife and myself that they want to be with us at every opportunity, even to sleeping in our pockets. They leap to us as a means of quick locomotion and for the pleasure of the leap and glide (pl. 4, fig. 2). Their behaviour has been treated at some length in the National Geographic Magazine for May 1947, “Flying Squirrels, Nature’s Gliders.” OBTAINING FOOD The obtaining of food is an ever-present problem for all animals. With some it is apparently a very simple process; with others it brings into play the use of the special structures with which nature has pro- vided them. The food-getting habits of practically every animal would make an interesting study, but only a few of the widely differ- ent methods can be mentioned here. The peculiarly shaped bill of the flamingo (Phoenicopterus), which looks as though it had been broken in the middle and the tip bent downward, is used in an interesting manner. As will be seen from the picture (pl. 7, fig. 1), the bird extends its neck almost straight downward, which brings the upper portion of the bill closest to the ground. The edge of the bill then scrapes along the ground in very shallow water, and the lower mandible opens and closes rapidly. The water thus scooped into the mandible is strained between the fine laminations of the upper and lower mandibles and then discharged after the minute crustaceans on which the flamingo feeds have been strained out. Relatives of the flamingo are the swans and other water fowl (Anatidae), many of which obtain their food in fairly shallow water. They eat a wide variety of plant and animal material such as seeds, green leaves, tubers, fleshy roots, mollusks, and crustaceans, and a few of them eat some fish. When the food is not too far below the surface of the water, they “‘tip up’ so that they can extend the neck and head to the maximum distance below the surface (pl. 7, fig. 2). Others dive for their food. Herons (Ardeidae) feed mainly on small fish, frogs, and other animal life, which they obtain by standing practically motionless in the water and waiting for the unwary victim to come within reach. They suddenly extend their long necks to a surprising distance and catch the prey before it has a chance to dart away. They may stand in such a position for hours at a time waiting for their victims to approach. ANIMAL BEHAVIOR—ERNEST P. WALKER 207 A very different method of obtaining food is employed by the birds of prey (Falconiformes), which catch their victims by a quick swoop. An outstanding example of a bird that employs this method is the falcon (Falco), which while soaring or flying almost out of sight will detect a bird in flight, or a bird or small mammal on the ground, dive from thousands of feet at speeds of as much as 280 miles an hour with the wings almost folded but extended just enough to enable it to steer itself, and strike and carry off its victim in its talons. In these dives the speed of the bird is so great that a whistling sound is produced. On one occasion I heard the whistling and an instant later a half-grown chicken was struck within about 8 feet of where I was standing, the falcon having come down at about a 45° angle over my head. It scarcely seems possible that a bird can fly with such speed and accuracy as to overtake and capture a startled duck in the air, but the duck hawk (Falco peregrinus anatum) does this regularly, providing a thrilling demonstration of skill and dexterity. Hunting with falcons was a royal sport in the Old World for many centuries and then was almost discontinued in Europe. It continued in Asia and is now popular again in Europe and the United States, where many different kinds of birds of prey are being trained to capture game or lures and bring them to the trainer. Among the birds that have been so trained are the beautiful, dainty little sparrow hawk (Falco sparverious), Cooper’s hawk (Accipiter cooperi), duck hawk, the prairie falcon (Falco mexicanus), golden eagle (Aquila chrysaetos), and even the barred owl (Strix varia). Another and little-known method of obtaining food.is practiced by the skimmers (Rynchops nigra), in which the lower mandible is con- siderably longer than the upper. Their regular practice in feeding is to fly so close over the water that the tip of the lower mandible is in the water, and when they come upon fish they merely lower them- selves enough so that they can scoop up the fish. Practically all woodpeckers (Picidae) feed on insects, most of which they capture in their burrows in wood. ‘These birds may often be seen going up or down trees, keeping the body often in a vertical position by grasping the trunk with the feet and leaning back on the stiff tail. Through their keen hearing they detect an insect under the bark or in the wood, and then proceed to cut away the bark or wood with their beaks until they approach close enough to the in- sect or worm so that they can draw it out of its burrow with their long-barbed tongues. The work of the woodpeckers in removing insects from the wood is far more beneficial than harmful, for if the insect is allowed to remain there it may do considerable damage, whereas the woodpecker ordinarily works only to excavate wood that has already been damaged by the insect, and by removing it prevents 278 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 further damage. The flickers (Colaptes), members of the woodpecker family, have found a less laborious method of obtaining a considerable portion of their food—they have learned to eat ants on the ground and have taken to feeding extensively on these and other insects in addition to what they get from the trees. The California wood- pecker (Balanosphyra formicivora baird2) and its relatives feed to some extent on acorns, which are very plentiful in many portions of their range, and they have adopted the practice of making holes in the bark of trees or in posts in which they place acorns for future use. Sometimes a trunk or limb of a tree or post will be studded ‘with hundreds of acorns, each in its own separate setting which holds it securely with the end of the nut projecting slightly from the hole. The night hawks (Chordeiles), whippoorwills (Antrostomus), swifts (Chaetura), swallows (Hirundo) and their relatives capture practically all their food in their very wide capacious mouths while flying. Some birds such as the tyrant flycatchers (Tyrannidae) and a few others have the habit of sitting on a perch that gives a good view of sur- rounding territory and watching for insects. As these come within range, they are captured in the air in a short flight by the bird, which then returns to its perch to watch for more. The hummingbirds (Trochilidae), as is well known, have a very long beak and tongue which they extend into flowers to feed on the nectar and insects that they find there. Other small birds known as honey creepers (Co- erebidae and Melithreptidae) have a long, sickle-shaped beak which they use in obtaining insects, nectar, and fruit pulp and juices. Feeding habits among the mammals also vary widely. The langur and colobus monkeys (Presbytis and Colobus) feed mainly on the leaves of certain trees. Many of the bats, particularly the small ones (Microchiroptera), feed almost exclusively on insects that they cap- ture in flight. Some forms are carnivorous, killing other bats and small birds and perhaps small mammals. Vampire and false vampire bats (Desmodus and Diaemus) lap blood that they obtain from warm- blooded animals by cutting off a very thin layer of the skin with their razorlike incisor teeth. Other very large bats with a wingspread of as much as 4 feet, which inhabit the Tropics, are known as fruit bats (Pteropus and related genera) because they feed largely on fruit. These bats frequently travel in considerable flocks and move to various regions from time to time to find food. Most rodents eat mainly plant food such as seeds or nuts, leaves, stems, or roots of plants; many, however, also eat some insects and small amounts of meat. A few have developed highly specialized feeding habits together with specialized teeth and shape of jaws. These include the fish-eating rat of South America (Icthyomys) and an insectivorous type (Rhynchomys) from Luzon Island in the Philip- pines. Nothing has been recorded regarding the habits of this animal, Smithsonian Report, 1949.—Ernest P. Walker PLATE 1 1. GOLDEN HAMSTER (MESOCRICETUS AURATUS) Lying on her back, she is trying to cut upward on the lower edge of a door. In this position she pulled herself through the %-inch crack under the door. 2. GOLDEN HAMSTER (MESOCRICETUS AURATUS) The cheek pouches are filled with seeds. The pouch is outlined by a white dotted line. ‘doip puv 00} 1940 oy} 9Ssvolor 07 ApBaI puv posBofar JOO} auO “VY SsII :19Y SUIP[OY o1¥ Jooj Jo soovjans szoddn ‘sso] Jo YYSuI| [[NJ 0} posaMo] ‘199 U90 ‘aspa ey} IOAO J[asioYy BursaMo] “WJay YOOTH SHL SACS SSAHON] OF ATSHS V WOY4 NMO”d ONILLAD (SNLVYENVY SNLADIMDOSAW) YSLSWVH N3G105 c 3a1lvid JI¥Je A “d IS2UIA—"Gp6| ‘Oday uetuosyztuISg Smithsonian Report, 1949.—Ernest P. Walker PLATE 3 1. EASTERN FLYING SQUIRREL (GLAUCOMYS VOLANS) Hanging head downward is one of its favorite positions while eating. 2. EASTERN FLYING SQUIRREL (GLAUCOMYS VOLANS) The flattened tail, long, strong fingers and toes, large eyes, and edge of the gliding membrane are clearly shown. Smithsonian Report, 1949.—Ernest P. Walker PLATE 4 1. EASTERN FLYING SQUIRREL (GLAUCOMYS VOLANS) A glide as seen from beneath and to one side. . 2. EASTERN FLYING SQUIRREL (GLAUCOMYS VOLANS) Preparing to alight from a glide. The two pictures on this plate were taken with the electronic flash which operates at about 1/5000 of a second. Smithsonian Report, 1949.—Ernest P. Walker PLATE 5 1. EGYPTIAN JERBOA (JACULUS JACULUS) A characteristic pose, with the very small arms and hands held close to the throat. The hairy pads beneath the feet are protection against hot sand and the abrasive action of rough surfaces. 2. TOWNSEND'S MOLE (SCAPANUS TOWNSENDII) (MUSEUM SKIN) Showing the broad, powerful forefeet with large, straight claws for digging; the lene, sensitive, mobile snout; short, sensitive tail; and dense, soft, short, plush- ike fur. Smithsonian Report, 1949.—Ernest P. Walker PLATE 6 1. THREE-BANDED ARMADILLO (TOLYPEUTES TRICINCTUS) Rolled up as a protection against enemies. The top of the head is the large inset portion in the front of the shell; the tail is lying along side of the head. (Photo- graph by Joao Moojen.) 2. THE BACK OF THE SAME ANIMAL The hinged arrangement of the plates of the back permit it to roll up completely. Smithsonian Report, 1949.—Ernest P. Walker PLATE 7 1. FLAMINGOES (PHOENICOPTERUS) The birds are shown feeding in shallow water, grooming, and resting. 2. WHISTLING SWAN (CYGNUS COLUMBIANUS) “Tipping up” to obtain food from the bottom. In this position these birds can reach to a depth of about 45 inches. (uslooyy ovor kq ydeisojoyg) ‘“xuey Joy uo sey puly yuo S}l puv ‘lapynoys Joy uo We 4YS syr ‘uIyO Joy Jopun pay SI YIIM JoyJOUL vy} OF Sursuryo st Aqeq W (SOLVNOYMOL SNHdOaVOS) HLOIS GANVYW 3uvuy SHL é ‘att 8 ALV1d £ a é ‘1oYYBJ 9yQ JO Yowuq oy} UO a1B SoIqed OMT, (SNHOOVF XIMHLIITNIVD) LESOWNYVYW GSELANL-ALIHM °1 oF JOHeA\ “q 1S2UIY—"GpG| ‘Oday ueruosy wg Smithsonian Report, 1949.—Ernest P. Walker PLATE 9 1. FEMALE SOUTH AMERICAN OPOSSUM (METACHIRUS NUDICAUDATUS) This animal lacks the marsupial pouch. Some of her babies clinging to her nipples are to be seen between her hind foot and her tail. (Photograph by Joao Moojen.) 2. FEMALE SOUTH AMERICAN OPOSSUM (METACHIRUS NUDICAUDATUS) Hight baby opossumsare clinging to the nipples between the hind legs of their mother. (Photograph by Joao Moojen.) "YOIS 94} JO pus oY} UO 4ooSUTI uv YO}Bd 0} YSnOUs Ivy pusejxo p[_nod snsuo0y ST {le} o[IsSuesyoid oy} puv qooy Suidsvis oy} 910N *ainyord SI Ul UMOYS [JOM ST JOO} “BLOUIBD OY} SuIyoyVM st ado YJo_ SYP =“ APQUepued oy} jo Aqrypiqe Suidseis o[qevyIvulel oy, “qySusy ut 492} 8 INOGB QUIT] 991} B SUTAIIVO ST 41 JOOF YU SII SAT UT -apul y1OM soo VSOYM piBzZIT poziperoeds ATYSIy VW (WIIWNd VHYNVSONDSIAN) NOAISWVHD “Z (iqa8v OSONOd) NVLNONVYO NVYLYWNS “1 Ol ALVId JAVA “d S2UIF—'Gp6] ‘qaoday ueiuosy qu ‘pavoq B 0JUO SUIPOY SI [rey S,4eyyou oy} Jo dry oy, ‘]I@] Joy JO oseq oy} punowe [1e} sit poddeam sey YOIM ‘Yyouq Joy UO SuIplA Aqvq B sBYy JouI0D jo] Joddn ayy Ul o[RUIEJ OY YT, (SNSOYSWNAA SAIALY) SABMNOW YaCIdS PM SBE Ace! JOWEA “d I89UIY— G6] ‘WOdoy ueruosyzrus Smithsonian Report, 1949.—Ernest P. Walker PLATE 12 aad te eae shoo Tere gree man tiara et PLAINS LEAST WEASEL (MUSTELA RIXOSA CAMPESTRIS) Upper and middle, in summer coat; lower, in winter coat (this form apparently does not become entirely white). This is a very rare form of which practically nothing has been recorded, and these may be the only pictures ever taken of it. About three-quarters natural size. “qSLIM 8 1OYING oy} UO SUTPFIS SI YOIYM (SUDI0a ShwwoznD)y)) Jolainbs sutAy udtojsvo uv IOAO SuIAy st yeq oy, re} opIsuoyoud A]5u0I4s St YIM JloszT SurApvoys st 4] (SnOsna SNOISA1ldy) LVG NMOYG Olg “Z (SNAW14 SOLOd) NOFYYNIM *1 €l 3LV1d JIA[EA\ ‘qq 1S9U14—"6} 6] ‘J40dey ueruosyAI US Smithsonian Report, 1949.—Ernest P. Walker PLATE 14 1. TWO-TOED SLOTH (CHLOEPUS DIDACSYLUS) Hanging beneath a limb is its usual pose. The forefeet have two toes, the hind feet, three toes. 2. YOUNG MALAYAN TAPIR (ACROCODIA INDICA) The striped baby coat is being replaced by the solid gray and black of the adult coat. Age 3 months. Smithsonian Report, 1949.—Ernest P. Walker PLATE 15 1. MALE BLACK AUSTRALIAN SWAN (CHENOPIS ATRATA) The bird is attacking a man in defense of the half-grown young in the foreground. The female was just to the left outside the picture. A AT UI A aii Lh PAAR RRO SIE iirc roe, ment es : «ee ae 1 wer ee : 7 Fe eo ; BEN he \ gn Js ol eee ion Sena F soe ease ee Be ae | 2. MALAYAN PORCUPINES (ACANTHION BRACHYURUM) The two animals alternate head to tail, in which position they regularly sleep. Smithsonian Report, 1949.—Ernest P. Walker PLATE 16 1. SIX-BANDED ARMADILLO (DASYPUS SEXCINCTUS) The animal is on its side asleep. They tremble almost constantly when sleeping. 2. THIRTEEN-LINED GROUND SQUIRRELS (CITELLUS TRIDECEMLINEATUS) ASLEEP Three rest on the top of their heads, one is lying on its back, and one in the back- ground is mainly right side up. ANIMAL BEHAVIOR—ERNEST P. WALKER 279 but we can judge from its structure that it probably eats insects or other soft animal food. The food of carnivores, the flesh eaters, is rather uniformly flesh or fish, but carnivorous animals frequently pass over the principal meaty portions of a carcass to drink the blood and eat parts of the viscera, thereby obtaining valuable vitamins. Almost invariably the glands or other parts which they eat are known to scientific workers as being excellent sources of vitamins. In one experiment that was conducted under my direction, we separated chicken viscera into its various component parts and then offered these to small carnivores. Almost all immediately chose the pancreas, which suggests that it is probably a valuable food. Most people have witnessed a dog burying a bone and later digging it up and chew- ing on it. It is possible that the bacteria of the soil act on the bone to make it more palatable and more digestible and possibly to elimin- ate any danger of ptomaine poisoning. A few carnivores, such as the binturong (Arctictis) and the palm civets (Paradozxurus and related genera), have taken to a diet consisting largely of fruit. An ingenious method was once demonstrated to me by the West African marsh civet or marsh mongoose (Atiliz paludinosus) in the National Zoological Park. This animal, about the size of a large cat, had a remarkable method of breaking bones. He was commonly given two pieces of horse ribs 4 inches in length and, when he had eaten most of the meat off of them, he would take a piece of bone between his forepaws, raise himself up on his hind feet with the hind legs well extended and with his forepaws well above the level of his head, and then quickly throw the bone down on the cement floor of the cage from a height of 2% to 3 feet. If he was not satisfied with the results of the first throw, he would repeat the process. The pro- cedure described suggests that these animals probably use the same methods in breaking the shells of mollusks and land crabs on which they feed in their native haunts of West Africa. The shortest mammal in the Americas and almost the smallest in the world is the lesser short-tailed shrew (Cryptotis parva) which inhabits the eastern United States. It weighs as much as two dimes, less than one-fourth of an ounce, and naturally such a tiny creature cannot cope with large antagonists in the usual manner. It normally lives in loose soil and leafmold where it feeds on earthworms, insects, and a wide variety of small animal life including frogs. Even a vigorous earthworm is a difficult creature for the tiny shrew to subdue in the usual manner, but earthworms lose their activity within a few seconds after the shrew gives them a few light bites; sometimes a single nip will suffice. Apparently the saliva carried into the very minor injury made by the shrew’s teeth is poisonous to the earthworms and takes effect very quickly. When the earthworm has become quiet the shrew proceeds to devour it, and it may be that this special- 866591—50-—_19 280 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 ized saliva accelerates digestion. Little is known regarding the poisonous bite of these little creatures, but it is apparently similar to that of a slightly larger close relative, the short-tailed shrew (Blarina) whose saliva is definitely known to be poisonous to many small creatures and whose bite on a man’s finger is sufficiently poisonous to produce pain extending halfway up the arm with resultant irritation lasting as much as a week. Such bites are somewhat com- parable to the bites of snakes in which a secretion from specialized glands is introduced under the skin of the victim, although these shrews do not have specialized teeth or glands that are primarily poison producers. Among the reptiles, the chamaeleon (Chamaeleonidae) has the interesting habit of obtaining the insects on which it feeds almost exclusively by suddenly extending its tongue to a distance almost as great as the length of its head and body and catching the unwary insect with it. The tongue is attached near the front of the mouth and folds back into the mouth like that of the frogs and toads, which also capture their prey in a similar manner (pl. 10, fig. 2). Emil Liers’ study of otters (Zutra) in North America have shown that these remarkably intelligent and playful animals feed largely on crayfish and numerous small invertebrates instead of fish, which they have generally been supposed to eat. It is strange that this was not discovered long ago, emphasizing the fact that there is still a fertile field for research on animal life. Through the long period of development of each species, animals have learned that in general each individual or pair requires a cer- tain amount of territory in which to sustain itself. Other species that do not conflict with it will be tolerated in this territory, while those that would be competitors may be driven out. Forms of life that constitute the food supply are rarely devoured to the point of extermination, and animals can therefore ordinarily obtain sufficient food within a rather definite territory. Some require but a small range for this purpose; others must have an extensive territory and move to various parts of it at frequent intervals to obtain the necessary food. For example, the wolf (Canis nubilus and related forms), an animal that preys on other animal life, usually has a range about 20 miles in diameter and, except when the female is living at the den caring for the young, it traverses this range, generally in large circuits, returning to any given portion of the range about once every 2 weeks. Seasonal fluctuation in the food supply, such as the migration of fishes or the dying off of insects, results in the migration of forms that prey upon them to locations in which food can be found. For example, insect-eating birds have an abundant supply of food in the Northern Hemisphere during the summer but, with the approach of winter, ANIMAL BEHAVIOR—ERNEST P. WALKER 281 insects and fruits become scarce and many of the birds cannot survive. By migrating to regions farther south, however, they are able to obtain an adequate supply of such food. Some forms that do not migrate change their feeding habits during the year. During the summer when fruit and insects and other animal life are abundant and easily obtained, they feed extensively on these elements, and in the winter they eat seeds and plant life that has matured during the summer. Good examples of this are the birds of the sparrow group (Fringillidae), in which the young are fed almost exclusively on insects and fruit and the parents eat such food extensively when it is available, but during the winter when few insects or fruits are to be had, they feed mainly on seeds. Others feed only on certain types of food and apparently cannot survive unless they can obtain these particular foods. Notable among these are the cross-bills (Loxia), birds of the Northern Hemisphere about the size of sparrows with usually some bronze, red, or purple markings. They feed on seeds of the spruce, hemlock, or pine which they obtain directly from the cones by perch- ing on them and reaching under the scales of the cones with their peculiar crossed mandibles and extracting the seeds. Such beaks are probably well adapted for this purpose but are a serious handicap to eating the type of food that most birds consume; therefore these birds are erratic in their occurrence in a given range. If there is a failure of the seed crop of the spruce, hemlock,-or pine, they must move to regions in which there is a good crop of seeds on these trees. Such sturdy, hardy animals as the American bison (Bison bison) and caribou (Rangifer) have extensive ranges that embrace both summer and winter forage grounds to which the animals migrate seasonally, some making a round trip of several hundred miles each year, measured in straight lines. We are prone to think of our food selection and handling as being superior to that of animals, but after observing the care with which animals select the choice morsels and pass over food that they detect as being contaminated or not palatable to them, I believe that the wild ones probably are far better fed and are better judges of food than we are. Studies of the feeding habits of antelopes on the native range disclose that in a 14-day period they eat 24 to 30 different kinds of vegetation. Possibly some of the material was eaten as many of us have taken food during war time, merely for the purpose of survival, but obviously they picked and selected foods that they preferred and thought were best for them. If one will watch a wild rabbit or almost any wild animal, he will see that it constantly and carefully selects or rejects food. On many occasions I have endeavored to induce animals, either wild or in captivity, to eat food that I had seen them pass over. Rarely could I get them to do so, and on one occasion, 282 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 the third time I offered a certain nut to a very friendly wild squirrel, it took the nut, opened it, and dropped it, perhaps to show me that the nut was not fit to eat. TRANSPORTATION OF OBJECTS The need of transporting food, nest materials, or young, and occa- sionally objects that apparently are taken for ornamental purposes or merely because the animal likes them, have been solved in various ways. Most animals carry objects in their mouths to some extent, just as a dog carries a stick or ball, but there are a number of ways in addition to this: for example, some of the Old World monkeys (Macaca and others) have cheek pouches that open inside the lips, but outside the jaws, in which they are able to store food, which must be a great convenience to them when they are trying to obtain their share in the presence of other greedy monkeys. They then grab what- ever food they can, put it in their cheek pouches, and eat it at their leisure. Monkeys with food in their pouches sometimes look as though they had mumps on one or both sides. Internal cheek pouches are found in a number of other animals such as the chipmunks (Tamas and Hutamias) and hamsters, which use them as shopping bags or baskets into which to place food to carry it home to the den where it is stored for future use (pl. 1, fig. 2). Another type of cheek pouch is the external one which is fur-lined. This is present only in a few North American forms such as the pocket gophers (Geomys, Thomomys, and related genera), the pocket mice (Perognathus) and kangaroo rats (Dipodomys and Microdipodops). Their pockets open outside the mouth, the skin of the face being folded inward to form the pockets. The pockets can be turned inside out to clean them, then pulled back into place by a special muscle. The owners of these pockets carry food, nest material, and even earth in them. We are so accustomed to seeing mother cats carry their kittens by grasping the skin at the back of the neck that we commonly think of this as being the principal way of carrying young; however, there are many other methods. The mother squirrel grasps the skin of the baby’s abdomen in her lips or teeth so that the little one hangs in an inverted position beneath her head, grasping her neck with its hands and feet and curling its tail around her neck, thus aiding in the carrying and leaving no dangling appendages to interfere with mother’s hands and feet in her travels through the trees. Baby monkeys cling tightly to the mother in most cases, but the white- tufted marmoset (Callithrix jacchus) mothers generally carry the babies only when nursing them; the rest of the time the fathers carry them (pl. 8, fig. 1). They cling to his long fur and ride on his back or under surface and hold on so securely that he can make leaps through the trees without dislodging them. Baby gibbons cling to the mother ANIMAL BEHAVIOR—ERNEST P. WALKER 283 on almost any part of her body and frequently take a position around her body almost like a belt. The Brazilian mammalogist, Joio Moojen, of the Museu Nacional, Brazil, has informed me that the baby of the rare sloth (Scaeophus torquatus) clings tightly to the mother and is almost completely hidden in her long loose hair. If no danger threatens, it may cling to her back, which is frequently the underside; however, if danger threatens, it is brought onto her chest or abdomen which is usually her upper side or is between her and a tree trunk. In this position it is well pro- tected between the mother and the trunk or branch to which she is clinging and is so well hidden in the fur that it can scarcely be seen (pl. 8, fig. 2). We commonly think of the marsupials as always carrying the young in the mother’s pouch. This is true for many of the forms, including the common opossums (Didelphis) of North and South America, but there are some marsupials that lack the pouch on the abdomen of the mother. In some of these species the young are carried or dragged about by the mother as they cling to her nipples. Good examples of this type are the small South American mouse opossums (Marmosa and allied forms) which range in size from about that of a house mouse to that of a common rat. The young hang suspended from the mother’s nipples which are in a cluster between her hind legs, and in this location they are well protected. When the young become larger, they are dragged on their backs, as the mother walks along the ground. If one becomes detached from the nipple, it is lost, for apparently they cannot again attach them- selves to the nipples, and the mother appears to make no effort to rescue them. The Metachirus pictured with the little ones hanging from her nipples had eight babies (pl. 9, fig. 2). Rodent babies of a number of different forms in various parts of the world also cling to their mother’s nipples until they are of good size, and when she travels about on the ground or through the trees, they dangle or are dragged about on their backs. The North American wood rat or pack rat (Neotoma) has this habit, and as many as four young, which individually may weigh almost a fourth as much as their mother, will be dragged about by her. In the Tasmanian “‘wolf”’ (Thylacinus), a marsupial, the pouch is a fold or shelf of skin on the abdomen between the hind legs which opens backward—a sort of rumble seat—and the young are carried in this. The best pouch of all is possessed by the kangaroos; they carry the babies in a large, deep, baglike fold of the skin of the mother’s abdomen until they may weigh almost one-fourth as much as the mother. When there are no young in this pocket, or when they are very little, it is shrunken and drawn up until it is rather small, but as the young grows the pouch stretches and can accommodate one 284 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 young until it is well able to take care of itself. There is a muscle around the entrance of this pouch which acts like a draw string and constricts the opening until it is very small, preventing foreign objects from getting in and the young from tumbling out. The very beautiful, graceful kangaroo rat mothers of North America lift their babies by grasping them by the back of the neck with their lips or teeth. They steady the little ones by holding them with the forearms, then hop along on their hind legs. This method of carrying the young was observed and photographed by that great naturalist, the late Vernon Bailey. The North American opossum is said to carry leaves and straw for its nest by wrapping the tip of its tail around the material. Some doubt exists, however, that this is an established habit, as only a few instances have been reported. As previously mentioned, gibbons, orangutangs and chimpanzees frequently carry objects grasped in their hind feet as well as in their hands. On one occasion I saw a chimpanzee pick up a piece of banana in one hand, a piece of bread in the other hand, and a head of lettuce with one foot. The ability to carry objects in the foot is particularly useful to animals that regularly traverse the forest by swinging by their arms from limb to limb. An orangutang carried a limb about 2% inches in diameter and 8 feet in length from an outside cage to an inside cage in the National Zoological Park, handling it much of the time by grasping it with the hind foot (pl. 10, fig. 1). A number of the animals that burrow, particularly those that make extensive tunnels such as the pocket gophers of North America, the mole-rats (Cryptomys.and Bathyergus) of Africa, and the bamboo-rats (Rhizomys and related genera) of Asia, push the earth before them, placing the hands close’to the sides of the head and against the earth and supplying the motive power for moving the body forward by the hind legs. Others scrape earth rearward with the forefeet and then send it farther rearward by strokes of the hind feet. This is the most common method of those that do not make continuous tunnels. The prairie dogs (Cynomys) of North America move a great deal of the earth that they use in building mounds by this method, and they also push earth before them. Beavers carry at least part of the earth or mud they use by holding it against the breast with the hands. Elephants (Elephas and Loxodontia) pick up objects by encircling them with the trunk. Apparently both whales (Cetacea) and seals (Pinnipedia) some- times hold their young to their sides by means of the flipper. This procedure has been observed so rarely and for such brief glimpses that little is known of it. Whalers say that a mother whale sometimes ANIMAL BEHAVIOR—ERNEST P. WALKER 285 uses this method to take her baby under water with her out of danger. Seals may do it for the same reason. Recently my wife and I witnessed a female black individual of the gray squirrel of the eastern United States collecting bark fiber for her nest. She went onto a dead limb of a tulip tree about 8 feet from our window and began loosening a strip of bark about a foot in length. Some was torn loose entirely but left hanging by a small strand, other parts were not hanging down but were well loosened from the branch. I thought at first she was not satisfied with the quality of the bark, and I was about ready to offer her advice on how to collect bark without wasting so much of it. Finally she began picking up the hanging fibers in her mouth, placing them crosswise and tucking the ends in so they would not drag and be in her way. When she came to the strands that still kept the pieces of bark attached to the limb she cut them off. In a few instances she backed up and pulled them loose by tugging. I then realized that she did not need any advice on how to gather bark with her equipment. She had loosened all the bark she needed but left it attached so it would not fall to the ground, then when she had enough loosened she began gathering it up. Of course she could not have had much loose bark in her mouth and still cut more bark loose from the limb with her teeth. As usual she had done her work in the most efficient way. Many times I have offered my pet “flying” squirrels a tidbit that they preferred to the one they were eating. ‘This confronted them with a real problem. Their experience through the ages has been not to drop food, for it falls to the ground and is ordinarily lost. They usually solve the problem by looking for a place in which to cache the food they are eating, then come back to me to take what I am offering. Likewise I have many times put the same problem or a similar one up to the gray squirrels (Sciwrus carolinensis) of the eastern United States. Most of them try to solve it by attempting to take the second morsel without dropping the first. Usually they cannot do this but make several attempts and finally finish by sitting nearby to eat the first piece and then come for the second, or sometimes they give up the attempt and take the first piece home or at some distance to eat, then return for the second. When I am giving them peanuts in the shells there is often an amusing struggle to get two together in the mouth so that they can both be held. Sometimes a squirrel will hug the nuts in its arms next to its neck and take a few short hops to get away and work on them at its leisure. I saw one old lady squirrel develop an ingenious method of solving the problem. After working with the nuts for a few seconds on several occasions she sat there and shelled one of the nuts, put the kernels in her mouth, then picked up the other nut quite easily. Later another learned this method and 286 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 both squirrels now do it regularly. Having learned the method, they usually hesitate after I have given them one nut to see if I will give them a second. Why make a trip home with only one nut when they can just as well carry two? The second squirrel to learn this is younger than the first one and may have observed the older one per- form the feat, for as pointed out elsewhere I believe animals learn much from observing others of their kind. However I saw the entire process of learning to do the act the first time with both of them, and thereafter it was a regular procedure to use the new accomplishment. Captive love birds (Agapornis) sometimes place straws under the upper tail coverts to transport them to the nesting site. My wife recently witnessed a novel method of transportation adopted by a yellowjacket (Vespula maculifrons). The bees were feeding on and carrying away pieces of raisins put on our fourth-floor window ledge for the birds. Generally they cut off a small piece of a raisin and never carried away a whole one, but she saw one bee roll a raisin with its head to the edge of the ledge and push it over. When the raisin started to fall, the bee followed it a short distance, then came back and repeated the process until it had dropped four raisins. My wife does not think the raisin was accidentally pushed over, as the bee’s movements seemed aimed at pushing it to the edge. Perhaps the nest was near the building and this bee had discovered an easy way of getting raisins to the nest. CARING FOR AND TEACHING THE YOUNG The type and amount of care and teaching that animal parents give the young varies from nothing to a very good education. Any consideration of this subject at once raises the question of how much the animal does by instinct and how much it learns from its parents or others. Of course, no conclusive or complete answer can be given to this, but there are many fragments of information that we can piece together to give us some light on the subject. Some animal mothers never see their young and give them no attention whatever. Among these are such animals as most snakes and lizards, which lay eggs that are hatched by the heat from the soil or from decaying vegetation, and the parent takes no part in their incubation. Exceptions are the pythons (Pythonidae) and skinks (Scincidae), which incubate their eggs, and the female alligator which stays near the nest to keep away intruders that might harm the eggs. However, the mothers take no part whatever in caring for or instructing the young. The mound- building birds of Australia and New Guinea (Megapodidae) do not incubate the eggs or care for the young; the mother lays the eggs in a mass of vegetation which she, together with other birds of the same species, scrape together, and in which all of them lay their eggs as in a sort of communal incubator. ANIMAL BEHAVIOR—WALKER 287 On the other hand many animals, particularly mammals and most birds, give a great deal of care to the young and obviously give them definite instructions. The bears (Ursus and Huarctos) are well-known examples. When mother bear begins leaving the den in the spring, the young are left inside and are apparently told to remain there. They do not begin coming out for some time—until the mother feels that they have developed enough to need a slightly larger world. She first permits them to play close to the entrance to the den while she stands guard, and later, when they are stronger, she takes them with her on foraging expeditions, at which time she tears open decayed stumps and logs to expose ants, grubs, mice, and other delicacies. Likewise, she turns over stones for animal food that can be found under them, digs up roots, and leads the babies where acorns can be found. Mother bear is a strict disciplinarian and does not permit the young to stray faraway. If danger threatens and she feels that for any reason she cannot take the young to the den, she often sends them up a tree while she stays on the ground not far away. She is usually successful in her instructions to them to keep silent, although occasionally a baby will become so frightened that it will ery, which often results in its being soundly spanked and cuffed for its infraction of her rules. If she desires not to send them up a tree but decides to run away, she keeps them close to her, and if they are disposed to lag or become tired, she will sometimes cuff them along ahead of her, sometimes tumbling them end over end so that they will have an incentive to keep up with her. Throughout this entire time, when the young are with the mother, observers have seen that it is definitely a training period in which the mother shows the young where and how to obtain food, what sounds and smells to avoid, and what are apparently safe in their haunts. Apparently about the same procedure is followed among the foxes (Vulpes), wolves and wild dogs (Canis), and wildcats (Felis), for the mother leaves the little ones in the den until they are able to begin playing about the entrance, where she finally permits them to sun themselves and romp and engage in tussles with brothers and sisters, gaining strength and agility. She brings them food, over which they struggle, and finally she brings live food so that they have the actual experience of handling live prey. Among the foxes and wolves, the father often participates in bringing food, and both mother and father stand guard near the entrance to give the alarm for the young to take refuge inside when danger threatens. Usually a short bark or two is sufficient to warn the little ones to take shelter. When they are old enough and strong enough to venture farther away from the den, the mother takes them on hunting expeditions on which they learn to catch small prey that is within their strength. On these expeditions 288 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 they learn to be alert, to beware of danger, to be on the watch for prey and how to catch it, and what to avoid. Mother deer (Odocoileus), antelope (Antilocapra), and moose (Alces) hide their newly born young and leave them hidden for some days. The baby antelopes may be left in plain sight on a grass-clad or very sparsely vegetated plain. Baby deer are hidden in the grass or sparse shrubbery, and baby moose may be hidden in such places or in slightly more dense vegetation. The mother then goes her way to get her food and rest, and returns to the little one only to nurse it at intervals of several hours. Thus the young are not exposed to the hazards of following the mother for the first few days until they have gained strength and are able to travel with such speed and endurance that they stand a good chance of survival by escaping with her. The mother cottontail rabbit (Sylvilagus) scratches out or selects a slight depression in the soil and lines it with fur that she plucks from the underside of her body. The depression is too small for her to be in it with the babies, but she crouches over it, and when the little ones nurse, they reach upward or climb up through the soft downy nest to reach her nipples. The cottontail nurses her little ones only at rather long in- tervals—apparently not at all during the daytime—and as long as 30 hours are known to elapse between feedings in some instances. When danger threatens she dashes away and the enemy usually follows her. Mother sea otters (Enhydra), which spend a great deal of time in the ocean, lie on their backs much of the time and the babies rest on the mother’s ventral surface. When she dives for food she leaves the little one floating on the surface while she goes to the bottom and picks up sea urchins and other food which she brings to the surface and eats while lying on her back. A mother flying squirrel that raised her family in my home, has given me many glimpses of how she cares for her babies. Flying squirrels are, of course, strictly nocturnal and there would be many hazards for them in the daytime; therefore I was not surprised to find that ‘‘Mother Glaucky” carries her babies back into the nest when she finds that they have strayed out during the daytime. Perhaps she and the tree squirrels teach their babies by demonstration and by voice, but I have not been able to detect much evidence of this. The young play among themselves about the nest, gradually gaining strength and agility and venturing farther from the nest. The length of time that baby flying squirrels and baby tree squirrels are weak and uncertain in their movements and are dependent on the mother is much greater than is generally supposed. Baby flying squirrels do not venture out of the nest until they are about 60 days of age and then only to explore in the immediate vicinity of the nest. By the seventieth day they are venturing somewhat farther, but their muscles are still soft and they have not gained agility or confidence. ANIMAL BEHAVIOR—ERNEST P. WALKER 289 In the wild they would probably not go farther than a few feet in their own nest tree. It is not until they are about 90 days old that they are ready to assume full activity. Golden hamsters, on the other hand, develop very rapidly, and by the thirty-fifth day, although not full size, they are apparently on their own in all respects. Swans (Cygnus, Chenopis, and Olor) are remarkably good parents and keep close watch that their young are protected from enemies. The female generally stays in the background and keeps the young with her while the male goes out to meet the intruder. If the danger is imminent he will approach with a direct rush and make vicious bites with his beak and will strike severe blows with his wings. If the danger is not imminent but he still feels an intruder might do harm he frequently approaches indirectly, that is, comes up to one side of the enemy apparently as though to catch it off guard. On one occasion while I was sitting very quietly on a rock in the swan yard trying to get a picture, the male swan persistently worked around to one side of me and when I did not move he finally grabbed my arm with his beak and tried to strike me with his wings. As soon as I started to move away he was satisfied. In this instance I had been trusting to my lack of motion to allay his suspicions, but my efforts to “‘freeze’”’ were not successful. One of the poses of the male black Australian swan (Chenopis atrata) threatening an intruder is shown in plate 15, figure 1. SLEEP Generally we think of sleep as a very simple state of inactivity which is similar in all animals, but actually the attitudes and types of sleep of various animals differ considerably. I think it likely that most, if not all, of the mammals that live in burrows well beneath the surface of the ground sleep very soundly, as they are comparatively free from danger while in their dens. Naturally, my observations of this fact have been limited and to my knowledge it has not been carefully studied; however, the few creatures of the burrowing type that I have been able to study all seem to sleep very soundly. A golden hamster can be picked up and handled gently within 30 seconds after it has ceased activity and has thrown itself down to go to sleep. I have similarly handled pocket mice when they were asleep in their nests and have found that they were quite difficult to arouse, which is in marked contrast to the great alertness of many other forms. Animals that live above ground are, of course, subject to a wide variety of hazards when they are asleep and must therefore, as it were, “sleep with one eye open.” This is particularly true of rabbits, most birds that sleep in the open and, no doubt, most other creatures that are in similar exposed locations. My pet flying squirrels selected a laundry bag hanging on a bathroom door for their nest and appear 290 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 not to be disturbed by the swinging of the bag, which perhaps simu- lates the swaying of the trees in which they would normally live. They are, however, distinctly disturbed by vibration produced by rubbing the door or the rod on which the bag hangs, which would probably remind them of the disturbance made by an enemy climbing to their nest in a tree. When thus awakened, they usually react in one of two different ways. If sharply startled they are very likely to dash out of the nest ready to take off in a glide or run to another, safer location; or if the alarm is not so sudden they may merely quickly and quietly go to the entrance of the nest and look out to see the cause of the disturbance. This behavior is, of course, associated with the rather exposed location in which they live, where danger may arrive from almost any direction and their best chance for survival may be to flee. I am inclined to think that the African elephant shrews (Macrosce- lides rufescens) sleep with both eyes open instead of only one. I have kept some in my den to study them at all hours of the day and night, and I have yet to see them with their eyes closed. They are invariably sitting upright with their eyes wide open, or at most only slightly closed. This trait suggests that they probably sit above ground in more or less exposed locations and are perpetually alert for danger. A wide variety of poses are assumed by animals in sleeping. In addition to the well-known attitudes of lying on the ventral surface, the back, or the side, a great many curl up in a very compact ball. The squirrels and others with bushy tails tuck the head and feet well inside and wrap the tail around them so that it actually affords some protection and warmth for the back of their necks and their backs. In this curled-up position they may lie on the side, but more frequently the head and feet are on the underside with the top of the head actually resting on the surfaces on which they are sleeping. This is a common position among a great many of the rodents. The giant anteater lies on its side, curls its head and feet together, and covers itself with its very long-haired tail which serves as a blanket and, perhaps in the wild, to some extent as camouflage. The sloths sleep hanging beneath a limb with the head thrown upward and forward so that it rests on the chest, or they may be partially sitting in the fork of a tree with the head forward between the upper arms, the tree trunk, and the chest. Some animals sleep standing up. Horses commonly do this, and some elephants do much of their sleeping standing. Most of the bats sleep hanging head downward, being suspended by the nails of their hind feet. The red bat (Nycteris), which sleeps hanging on a twig in a tree, has an extensive membrane between the hind legs which it draws downward so that it serves as a protection to the ventral parts of the body. When hanging head downward, ANIMAL BEHAVIOR—ERNEST P. WALKER 291 bats are in a good position to take off for flight, for they are generally at an elevated location and have merely to let go with their toes and spread their wings to be in full flight. I have noticed that the nine-banded, the six-banded, and the hairy armadillos (Dasypus and Huphractus) all tremble almost continuously in their sleep, particularly when lying on their backs or sides, as they often do. This is unique among mammals with which I am ac- quainted, but I have no theory to explain it. Malayan porcupines (Acanthion brachyurum) like to sleep side by side and have an interesting method of avoiding the spines of another that has already lain down. Each succeeding one merely faces in the opposite direction from the last one in the row. I have seen as many as five lying asleep in such alternating head and tail positions, but when I tried to take their pictures in this arrangement I usually disturbed some of them, so that I have never been able to photograph more than two together (pl. 15, fig. 2). The tiny bat parakeets (Loriculus) sleep hanging head downward, clinging to the perch by their feet. Of all the small mammals I have observed the females are much more particular regarding the nest than the males. The females will move it about, cut on the nest box, assemble nest material and keep it well shaped into a snug nest, whereas most of the males are far less particular, usually working on the nest only enough for it to be passably comfortable. Of course, my wife noticed this before I did and pointed out that females have stronger instincts for home main- tenance than males. Most mammals, when they have the opportunity to awaken natur- ally, like to sit and “think,” or perhaps just sit, like many people who cannot start off ‘in high gear.’”’ Those that I have observed, after about half an hour start normal activity. Hibernation was discussed briefly in my previous paper on Animal Behavior and has been extensively treated in other literature so will not be further mentioned herein. CONCLUSION The better our knowledge of animal habits and behavior, the better we are prepared to cope with the problems in connection with animal life and the administration of resources in which animals play a part of greater or less importance. Biologists are constantly being con- fronted with problems of how to control, circumvent, keep away, or increase wild life. The problems may be simple or very complex, but are always interesting. Wildlife administration has become an important branch of national, State, and local government work for we have come to realize that many of the forms are highly beneficial and should be protected and 292 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 built up to the maximum possible numbers, and that a few are injurious and should be controlled. As the activities of mankind are extended, the importance of wildlife protection increases correspondingly, for we could not live without animal life and the extirmination of any form is a serious loss. Better understanding of animals leads to recognition of their value and there- fore to more interest in their protection, and the study of animal life as a profession, as a hobby, or merely through casual observation yields much pleasure. Norr.—All photographs are by Ernest P. Walker unless otherwise listed. THE BREEDING HABITS OF THE WEAVERBIRDS A STUDY IN THE BIOLOGY OF BEHAVIOR PATTERNS By HERBERT FRIEDMANN Curator, Division of Birds, U. S. National Museum [With 8 plates] The weaverbirds, as their popular name implies, are, by and large birds noted for the elaborate nests they build—in many cases actually weave—out of grasses, straws, rootlets, and other similar materials. Included in this family (Ploceidae) are some of the finest, most expert, and most famous of all avian architects. In no other single bird group of similar status has the habit of nest building been carried to greater heights, indeed their only rivals for excellence in this particular are some of the hang-nests or troupials of the New World. My interest in the weaverbirds began some 30 years ago when I had the opportunity of studying the actual weaving methods in captivity of one species, the red-billed weaver, Quelea quelea. Not long after- ward when I began my studies of parasitic birds I learned that at least two species of weavers, and possibly several others, not only built no nests at all, but laid their eggs in the occupied nests of other kinds of birds to whose care the eggs and the subsequent young were left. A few years later, over a year’s field work in South and East Africa gave me ample opportunity to become familiar with the surpris- ing range of aspects the nest-building habit exhibits in this family of birds. In this paper it is not my purpose to attempt to describe each and every one of these aspects, but to discuss them from the point of view of the biological implications they present. ‘The family is a large and varied one and contains a great many species (about 275), many of which have highly divergent nesting habits, which offer very suggestive data to the naturalist concerned with the evolution of habits in birds. In order to appreciate more adequately the significance of the various aspects of the nest-building habit in these birds, and to evaluate them more properly in connection with other parts of the life histories of birds involved, it is necessary to make a few preliminary 293 294 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 general remarks. Birds, as a rule (there are, of course, exceptions), go through a fairly definite cycle of behavior patterns during the year, and repeat this cycle every year of their adult lives. Very briefly stated, the cycle consists, in typical fashion, of the following parts: Migration or, in the case of nonmigratory birds such as the weavers, the fragmentation, or breaking-up, of “winter,” nonbreeding flocks into pairs of birds; the establishment of individual breeding territories (the extent and definiteness of which vary in different species) ; court- ship and mating; nest building; egg laying; incubation; care of the young; and, finally, migration or the return of the individual birds or pairs to the “winter” flocks. Each of these parts of the whole cycle is subject to great variation, and each may be influenced in its devel- opment and expression by its antecedent stages, and each may exert a similar influence on its succeeding stages. Like any other sequence of events or stages, the undue development of any one part may tend to throw out of line one or more of the remaining parts. When everything runs smoothly according to what may be looked upon as a normal pattern, it is very difficult to observe the sequential effects of each part on its successor and it is rarely possible to get even vague glimpses or hints of how the per- fected whole cycle came to be developed through the ages. It is only when something deviates from this normal pattern that we have much chance of learning anything of the factors that control or influence it. With these general thoughts in mind, it is instructive to examine in some detail what has transpired in one family of birds with respect to one part of the life cycle, the nest-building habit, and, in connection with it, to related portions of the cycle. The weaverbirds have been divided into a number of subfamilies (the exact number differing in different classifications), which in a general way are characterized by different nest-building habits. The arboreal, so-called typical weavers (Ploceinae) construct a suspended type of woven nest, usually shaped like a ball or a closed oval with a lateral or a downward-extending entrance tube or “vestibule.” Most of the species of this group are colonial, some- times as many as 50 or more nests being built in the same tree, and often with no others on any of the surrounding trees for considerable distances. On the other hand, some species are quite solitary, like Reichenow’s weaver (Ploceus reichenowi), of which usually only one, and apparently never more than two pairs nest on the same tree, and in those cases where there are two, only one seems to be breeding at atime. The instinct to build is, however, very strong in most members of this group as farasknown. Thus, Bates (1930, p. 484) noted of the hooded weaver (Ploceus cucullatus) in Cameroon that ‘. . . it seems to be a necessity of the birds’ nature to be always busy with their nest; they will occupy themselves in their spare time with tearing down BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 295 unused nests, strewing the debris on the ground under the tree . . .” Of the spectacled weaver of South Africa (Ploceus ocularius) Roberts (1940, pp. 337-338) records that the nest has a protruding entrance tube which is usually from 3 to 6 inches in length, but that in several instances the nest-building activities of the birds were so great that the entrance tubes became extended to a length of 6 feet! One nest in the Albany Museum at Grahamstown, recorded by Stark, has an entrance ‘‘upwards of 8 feet long.” It is a well-known occur- rence in captivity, such as in zoologicalparks, for different species of weavers, when given quantities of raffia or straw, to weave a vast quantity of rather formless masses of compact, densely woven mate- rial over the branches in the cage and even over the wire mesh of the cage itself. The birds do so equally avidly whether they are in breeding or in nonbreeding plumage; in other words, the urge to build, which in most birds is seasonal and is part of the cyclical sequence of behavior patterns, is here extended far beyond its normal limits. Furthermore, in at least some of the species of typical weavers (many are still very poorly known as far as details of habits are concerned), the bulk, if not all, of the actual nest construction is done by the males and not by the females. (In most ordinary birds the female does most of the nest building.) Thus, in the masked weaver (Ploceus velatus mariquensis) Taylor (1946, pp. 145-155) found that the males did all the nest building, except for some of the lining which was put in by the females. When a nest is completed the male that built it immediately starts to make another, and in one colony a single male wove no fewer than 11 completed nests. In another species, the Baya weaver of India (Ploceus philippinus), Ali (1931) found that the males were polygamous and that the number of mates each was able to get depended on the number of completed nests he was able to build for them, the actual courtship and mating behavior taking place in or around the newly finished nest. Ali writes that in his experience with this species— ... in the initial stages of an adult nesting colony, no hens are as a rule in evidence, and I have been unable to discover their whereabouts during the first few days. It would appear that the instinct to breed asserts itself earlier in the adult cocks than in the hens, for it is not until the time when the nests have progressed to a stage where the egg-chamber is finished or nearly so, that some of the females first become physiologically “‘ripe.”” They now visit the colony quite obviously with the sole object of “‘prospecting”’ for laying sites, i. e., to discover if there are any nests that are ready for their occupation. ... Two hens often fight for the possession of an acceptable nest. The successful hen henceforward boldly enters the nest and busies herself with finishing off and making the interior comfortable. In no case have I been able to observe the cooperation between male and female so often described. The lion’s share of the building—in fact all of it—is undoubt- edly done by the cock alone. Her contribution is only the “‘interior decoration”’.. . The “building mania,’ as it has been called, that comes over the adult cock at this season is a sure indication of his readiness to breed... 866591—50——20 296 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 While my own field notes are less detailed for any one species of weaver than are Ali’s on the Baya, I found males of a dozen or more related African species actively engaged in nest building. In the cases where my notes are fullest (the Cape weaver, Ploceus capensis olivaceus, the black-headed weaver, Ploceus nigriceps nigriceps, and the Kenya vitelline weaver, Ploceus witellinus uluensis) my observa- tions indicate that the male does most of the nest weaving, if not all of it. In some other species (Jackson’s weaver, Ploceus jacksoni, and the spotted-backed weaver, Ploceus spilonotus) there seemed to be more activity among the females in this regard but still the bulk of the building was carried on by the males. (It may be that some of the “females” were really males in nonbreeding plumage, a point that could have been determined only by more collecting at the time. That this is not unlikely may be adduced from the fact that in my field notes on the masked weaver, Ploceus velatus arundinarius, I wrote that the females take part in the nest-building activities, but Taylor’s careful study of a slightly different subspecies of this bird, referred to above, convinced him that the males did all the actual construction and that the females merely added or rearranged some lining materials. It may well be that the birds I recorded as females were males that had not yet come into nuptial plumage.) Chestnut weavers (Ploceus rubiginosus), watched in captivity, showed more nest-building activity among males than females. Of Speke’s weaver (Ploceus spekei) van Someren (1916, p. 409) noted that, ‘‘dozens of nests are built by the male, but only one is occupied; thus there are always plenty of old nests in all stages of completion.” In one of the forest-dwelling, relatively solitary, or at least not highly colonial, typical weavers, Malimbus cassini, of West Africa, Bates (p. 478) found that both sexes take care of the young. He shot a male and a female at their nest, which had, ‘‘. .. a woven entrance tube 2 or 3 feet long, so thin that its walls were transparent, and the birds could be seen entering and leaving, feeding young.’’ In the Cape weaver, Ploceus olivaceus capensis, previously mentioned, Skead found that both parents fed the young, and it appears that this behavior is fairly widespread in the entire group. Another section of this subfamily contains the so-called bishop birds (Huplectes) and the whydahs (Coliuspasser and its close allies). These birds are far more terrestrial than the members of the genus Ploceus and their habits are somewhat different. Lack (1935, pp. 817 ff.) studied the fire-crowned bishop (Huplectes hordacea hordacea) in Tanganyika Territory and found that the males are polygamous and do most of the nest building, each female merely finishing or lining the one it occupies, and each hen continuing to add to the nest throughout the period of incubation, eventually making it so thick that the ob- server can no longer see through it, and adding a small saclike ledge BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 297 above the entrance. One male was found to have three mates at the same time, and may have had still others. The males have very definite territories and the hens apparently seek out the established cock birds. Actual mating takes place when the nest is in process of being built by the male, but the female has, in all probability, already settled in his territory for some time before this. Incubation of the eggs and feeding of the young are solely the business of the hens, which incubate chiefly at night, the warmth from the sun presumably being sufficient for the eggs during the daylight hours. Less complete data on the whydahs (Coliuspasser ardens and C. jackson) suggest that these birds are monogamous. Thus, in writing of the latter species Jackson (1938, pp. 1469-1470) states that he re- gards the evidence against the supposed pologamy of this whydah as conclusive. Near Nairobi, Kenya Colony, he had extremely favor- able conditions for watching this species within a fenced-in enclosure with open grass, outside of which enclosure the grass had been burned. In such a small area it was easy enough to count and mark with a stick each dancing ground. This done, the whole area was hurriedly quartered with the aid of two boys to move and frighten away all the birds present; we then retired a short distance, sat down and waited for them to return. The cocks very soon appeared; the females were much more wary, but returned in due course. Some of them settled in the grass and remained there, evidently on their nests, while others were occupied in going to and fro with fine grass in their bills; these latter rarely remained hidden for more than a minute at atime. Next day we returned, and by quartering every yard of the area we discovered four nests with three eggs, three with two eggs, one with a single egg and three not yet completed. Each nest when found was marked by tying a knot in a wisp of tall grass close by. At the end of a week we again returned; but no more nests were found, and on no occasion did the females equal the number of cocks, but I accounted for this through my failing to detect one or two of them as they sneaked back to their nests containing incubated eggs .. . At least it seems from this account that there were not more hens than cocks as would have to be the case in a polygamous species. Unlike many of the weavers whose courtship is performed at the newly constructed nest, a group of males of Jackson’s whydah makes a roughly circular dancing area by breaking or snipping off the tall grass to make a clearing of from 2 to 6 feet in diameter. In the middle of this clearing are left standing a sizable number of grasses forming a dense tuft into which the males partly excavate little re- cesses. Then a number of males go through a leaping peformance, generally with no hens around to watch them. Jackson describes the position assumed by the cock as follows: The head is thrown back like that of a proud Turkey-cock, the beak being held horizontally; the feet hang downward, the tail is held straight up till it touches the ruff at the base of the head and neck, the ends of the feathers falling in a curve downward, with the exception of two tail-feathers which are held straight out and downward. 298 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 While actually rising in the air the half-open wings are worked with a very quick shivering motion, and the feet are also moved up and down very rapidly, beating the air. The bird springs straight up in the air, sometimes for a few inches and sometimes to the height of two or more feet, and then drops. The whole of the plumage is much puffed out throughout the performance, which is repeated five or six times, with a short interval for rest. The game would appear to be somewhat fatiguing, as the bird rarely makes more than five or six jumps at a time without a short rest . . . They very often assume their curious jumping attitude some little distance before they arrive at their playground. . . Besides the data quoted above it may be added that apparently but one male may make use of one dancing area, and that often at the end of the jumping dance it appears to try to burrow into the shallow recesses of the central tuft of grasses (as though there were nests there). Jackson’s whydah shows the courtship behavior pattern developed to a greater degree of display and ostensible rivalry between males, and to a greater specialized areal usage (courtship, or dancing, ground as contrasted with nesting site or even breeding territory) than others of its relatives, but the difference seems to be more one of degree than of kind. As typified by its habits, this section of the terrestrial Plo- ceinae may be said to be characterized by elaboration of courtship behavior from individual displays near the nest to a complex display at a dancing ground, and by what seems to be monogamy. (More detailed information is needed on this point, however. In the case of the black whydah, Coliuspasser concolor, and of the red-collared why- dah, Coliuspasser ardens, there are observational data supporting a monogamous state; in the long-tailed wydah, Coliuspasser procne, similar but less extensive data suggest polygamy.) Recently V. D. van Someren (1945, pp. 131-141) has concluded that Jackson’s why- dah is polygamous, but his own presentation of the case is not too pos- itive. Thus, he writes that— . polygamy appears to be general, and seems to arise because of the imper- fect correlation between the maturation of the males and females. Some males mature early, others late, and the early males may cease dancing and start moult- ing while later males are just beginning to assume breeding plumage and dance. This irregular maturation of the males may be spread over several months, while by contrast, the females mature almost simultaneously, and all nests are found at the same stage of building or incubation within a few days. Since the sex- ratio of the mixed flock is almost 50:50, late maturing males are thus able to mate with several females, because the mature females probably now outnumber the mature males. Males may commence dancing some four months before the first nest is found, but these early males are probably unsuccessful at mating be- cause of the unready state of the females. Males may start dancing while still in non-breeding plumage, but the behaviour pattern of these immature males is undeveloped in several respects. BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 299 It may be noted that the assumption that late-maturing males are able to have several mates is based on the thought that more females are mature in the later than in the earlier part of the season, but, only a few lines above, van Someren informs us that all the females mature simultaneously, which would imply that the females mature later than the early-maturing males. It would seem, from this, if van Someren’s assumption be correct, that the early-maturing males might be no longer in breeding condition late in the season when mates are available, or else they would compete with the later-maturing cocks for the hens to a degree sufficient to diminish the likelihood of the late-maturing mates having more than a single mate apiece. The care of the eggs and the young appears to be left wholly to the females, and, as far as the incomplete evidence goes, the actual nest building is also done by the hens of the various species of whydahs. We have but little reliable data as to the territorial aspects of the lives of most of these birds, except for Jackson’s whydah. In his study of this bird van Someren found that— . . . true territorial behavior becomes evident early in the sexual break-up of the flock. The males, isolating themselves on rings (7. e., dancing areas) establish a well-defined territory of small extent, of which the ring itself is the focal point; the territory extends all round the ring at a radius of 6 to 10 feet from the central tuft. A female alighting anywhere within this territory may be solicited by courtship behavior by the male on the ring, even though she may not alight on the ring itself. Another full-plumaged male alighting on this territory is treated in one of two ways, depending on the attitude of his tail as he alights. If he alights with his tail arched and the two outer plumes drooping as in the dancing attitude, he is attacked with pursuit flight if the owner is present in the territory. If however, the intruding male alights with his tail folded in the normal flight attitude he is usually solicited and displayed to be the owner as if he was a female. It is very noticeable that when a male returns to his territory from outside it, the tail is arched and the two outer plumes drooped the moment he crosses the boundary; the bird alights in the dancing attitude, and thus shows his ownership by his appearance... Where two or more rings are found within a few inches of one another... they are all formed by the one male, who may use them alternately while dancing, and keep them all in good order . . . rings occupied by two separate males are not found closer than about 12 feet. These boundaries are accepted by the other members of the flock early in the break-up, hence territorial squabbles are seldom seen late in the season . . . This territory is related purely to sexual functions and has no food significance; feeding is carried out in a mixed flock even in the height of the dancing season, on neutral ground where sexual rivalry is notably absent. Furthermore, this territory appears of no significance to the females apart from the fact that they are attracted to the rings; they are unaware of the boundaries of the male territories. At nesting time, the males cease dancing vigorously and the main dancing area may become completely deserted; the females nest in a different area which is usually some distance away from the dancing grounds. The 300 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 nests tend to be grouped together, and are usually about 20 to 30 feet away from the nearest ring if males have been dancing previously in the neighborhood, i.e. well outside the territory boundaries. In connection with the development of courtship posturings, it may be pointed out that, unlike the members of the arboreal Ploceinae, the whydahs, and, to a somewhat lesser extent, the bishop weavers, have very marked sexual dimorphism in the breeding plumage of the adults and many species have elongated or otherwise specialized plumes in the nuptial dress of the males. In the nonbreeding season the sexes generally look alike. Before going on to the next subfamily, we may briefly summarize the situation in the typical weavers (Ploceinae). Many of the species are colonial (which means, in effect, that in most of them the individual nesting territories are nonexistent), and in those species that have been most fully observed the nests are built largely or wholly by the males. Furthermore, in some forms it seems that the males are regularly polygamous and that the number of mates each one acquires depends on the number of nests he has been able to provide for his mates. From the standpoint of our hypothetical “standard” picture of the cycle of breeding behavior patterns this implies that the first stage— migration or the fragmentation of flocks into individuals or pairs—is omitted, that territoriality is likewise skipped, and that the usual sequence of courtship and nest building is reversed. As we have noted, the actual courtship and mating takes place in and about the newly completed nests the males have constructed. The Ploceinae take 2 years to acquire adult plumage and to come into breeding condition. This summary is correct as far as it goes, but there is still one more variation in the reproductive behavior pattern exhibited in this sub- family. One species, sometimes called Rendall’s seed-eater, some- times (and more properly) the cuckoo finch, Anomalospiza wmberbis, a bird with no very close relatives, but apparently nearer to the bishop weavers than to any other assemblage, is wholly devoid of any nest- building or incubating or rearing instincts, and is, in short, parasitic. It is still very imperfectly known, and all that may be said with any certainty is that it is parasitic on small ground-nesting (or near the ground nesting) warblers of the genera Cisticola and Prinia. (Six species of the former and one, possibly two, of the latter genus are known as hosts of the cuckoo finch.) As Delacour (19438, p. 71) has recently pointed out, the fact that Anomalospiza agrees with the Viduinae in being parasitic does not necessarily imply close relation- ship to the members of the latter group. In most respects it seems best placed, taxonomically, with the Ploceinae, but, it must be ad- mitted, is an aberrant member of that group. It is aslightly gregari- ous bird, living, at least in the nonbreeding season, in loose flocks. Nothing is on record concerning its courtship, sexual relations, or territorial habits. BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 301 The next subfamily is a much smaller assemblage—the buffalo weavers (Bubalornithinae)—containing only two species, each with several races. Unlike the typical weavers these birds do not construct nests of fine weaving and elaborate structure, but make rather bulky, massive nests of twigs and thorny branches, rough and untidy in general appearance, and not suspended from, but placed on top of, the branches of large trees (in my experience often baobabs). The birds are colonial and the nests are often placed so close together that they are actually in direct contact with one another. Jackson (p. 1380) records that he saw where branches had given way under the combined weight of too many of these nests on several occasions. Brehm found as many as 18 nests in 1 mimosa tree in northeastern Africa. The nests are often 2 or 3 feet across and are used and repaired and added to year after year. Each nest of the black buffalo weaver (Bubalornis albirostris) contains two or more chambers, lined with grass and straw, each with an entrance facing away from the other. The only nests I ever saw of the other species, the white- headed buffalo weaver (Dinemellia dinemelli), had but a single cham- ber. There are descriptions in the literature of nests of Bubalornis containing more than two entrances. Priest (vol. 4, p. 220) writes of one that “‘. . . there were numerous entrance holes, and it looked as if about a dozen birds lived in each of these communal nests . . .” That the urge to build is extended in these weavers beyond the usual small part of the annual cycle of most birds, as it is in the Ploceinae, is indicated by some observations made in Darfur, in the Sudan, by Lynes, who noted that “‘. . . at all seasons we found these Textors (=Bubalornis) hanging about their everlasting great nest-clusters, into which, even in midwinter, birds with quite inactive sexual organs would sometimes carry twigs as if nesting . . .” Many years ago, in southwestern Africa, Andersson described the nests as follows: The collective nests consist externally of an immense mass of dry twigs and sticks, in which are to be found from four to six separate nests or holes of an oval form, composed of grass only, but united to each other by intricate masses of sticks, defying the ingress of any intruder except a small snake. In each of these separate holes are laid three or four eggs . . . I obtained no less than forty of these eggs . . . and on the following day the birds were busy in repairing one of the collective nests which had been injured during the collection of the eggs .. . I believe these nests are annually added to, for, so far as I have been able to see, the same nest is retained for several consecutive seasons. We do not have nearly as complete information on the buffalo weavers as on the typical ones, but what data are available indicate that the birds are colonial, that there is little or no observable evidence of any prenuptial fragmentation of wintering flocks, and that the males do at least part of the nest building. Whether they do it all or not is still unknown. 302 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The evidence does suggest that the nest-building habit is indulged in out of the breeding season and by nonbreeding birds; not too different from the typical weavers in this respect. The next subfamily, the sparrow weavers and social weavers (Ploce- passerinae), shows a great indulgence in nest building, in and out of the breeding time, which culminates in the truly gigantic communal structures of the social weaver (Philetairus socius). Thus, of one South African sparrow weaver, Plocepasser mahali, Roberts (1940, p. 332) writes that— . a single pair of birds will construct as many as a dozen nests of whitish grass stems on the projecting branches, these being arched over the top with two entrances below on opposite sides, so that there is no cavity for the eggs and evidently made for amusement only; the nests in which the eggs are laid have only one entrance and are warmly lined with feathery grass tops .. . A somewhat different description is given by Stark (1900, pp. 84-85) who informs us that the species is— . of social habits, it remains in flock all the year round and breeds in company, several nests being generally built in a single tree. Rarely have I met with more pugnacious birds; the males in spring are constantly fighting, and so desperate are their quarrels that the combatants frequently lie exhausted, side by side, on the ground, incapable for further movement ... The nests are large, roughly built, kidney-shaped structures, usually placed near the ends of the branches of a mimosa or other thorny tree. They are constructed of long grass-stems, the blades and flowering tops being woven together, the stiff stalks projecting in all directions. During the winter each nest has two entrances from below, separated in the interior by a narrow bridge of grass, on which the birds roost. At the beginning of the breeding season one entrance is stopped up with leaves and grass, a shallow cavity being left in which the female deposits two or three eggs... As soon as the young are on the wing, the second entrance is unstopped, and the nest is again used, both by the old and young birds, as a roosting place. The nests are annually repaired and last for many years. A somewhat similar account holds for another species, the gray- headed social weaver of East Africa, Pseudonigrita arnaudi. Jackson (p. 1384) comments on the remarkable nest of this bird— . . . it is very large and quite exceptionally compact, and has two entrances pointing downwards. During the period between breeding seasons these nests are used for roosting, the birds resting on the ridge between the two entrances. In the breeding season one of the holes is stopped up and the eggs are deposited in a depression beyond the ridge on the material used for stopping up the second entrance. The nest is firmly woven to several twigs or branches . . . in small clumps of five to eight nests together. Of a slightly different race of the same species Jackson found (p. 1385) the nests ‘‘. . . were packed together so closely as to form almost one compound nest.” Probably the most remarkable of all weaverbirds’ nests is that of the famous sociable weaver of the western arid portions of South Africa, Philetairus socius. The enormous communal nests built by BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 303 these little sparrowlike birds attain truly great proportions—as much as 25 feet long and 15 feet wide at the base and from 5 to 10 feet in height! While each nest is the product of not a lone individual, or even a pair, but of a whole flock of as many as 75 or 80 pairs, still the sheer bulk of the nesting material gathered and placed by the birds is a striking testimony to the tremendous year-round urge of the nest- building instinct in this species. The time I spent studying this bird in the western Transvaal in 1925 was one of the most fascinating experiences an ornithologist could have, and I cannot refrain from including here part of my notes, at the expense of repeating some items already published in an earlier paper (1930). As the common name of the bird implies, Philetfairus is very social in its habits; in fact it is probably as social as any bird could possibly be. It is always found in flocks, feeds in flocks, and breeds in large, many-apartmented compound nests. The smallest flocks that I saw contained about 20 birds; the largest one at least 150. The flocks seem to stay pretty much in the same general vicinity all the year round, and the birds use their huge, massive nests as roosting places during the nonbreeding season. With this extreme sociability and sedentary habit of life the territorial relations of the species have-been modified in a way that is quite remarkable, perhaps unique among birds. Instead of each pair of birds having its own breeding territory, each flock seems to have a definite territory, and as the individual flocks are usually far enough apart not to compete with each other, the boundaries of these territories are seldom crossed by individuals of other flocks and other territories. However, in a few cases in my own experience two flocks were fairly close together (i. e., two nests were on trees not very far apart), and the birds mingled more or less while feeding, but in these cases far more fighting and quarreling was observed than in all the others together. In an area approximately 100 miles long and 10 miles wide, or 1,000 square miles in all, I found only 26 nests of the social weaver, so it can be seen that the flocks ordinarily do not live in very close juxtaposition to each other. (The nests are so large, and so conspicuous at great distances, and the country so open and easy to examine, the trees being so relatively few in number, that I am quite certain I found practically every nest in this area.) The nests observed varied in size as did the flocks. The smallest nest found measured about 3 feet in diameter at the base and was about 3 feet high and had about 10 entrances on the under surface, indicating that it contained that number of individual nests. The largest one found was incomplete, i. e., a piece of it had broken off, breaking its supporting branches by its weight, but the remaining part was a large, flat, horizontal mass of straw, more or less repaired at its broken edge, and measured about 25 by 15 feet at the base and 304. ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 was about 5 feet high. The part that had broken off must have been about 5 feet in diameter each way. This nest contained about 95 nests within it. In a locality where these birds occur it is impossible to remain long unaware of their presence. ‘Trees are not so numerous but that each one becomes an object of more or less importance in the landscape. Needless to say a tree on which there is a social weaver’s nest is a very conspicuous object, visible for a great distance and widely proclaims the presence of the builders. But the birds themselves soon intrude upon one’s consciousness with their noisy, harsh, chattering notes as they fly by in flocks or feed in scattered bunches upon the seeds of the small, stunted shrubs and plants that wrest an existence from the inhospitable soil. While feeding they keep up an incessant chatter much like a flock of house sparrows, and, like them, frequently quarrel over bits of food. In flight they all act in unison with a precision quite remarkable for birds of their type, the whole flock turning, rising, falling, wheeling, and stopping more or less together. Although the birds live in compound “‘apartment-house’’ nests, feed and fly in flocks, and are at all times exceedingly gregarious, they seem to establish fairly strong mating relations as far as my field observations indicate. If they were haphazardly promiscuous they would be forever in each other’s way getting in and out of the entrance holes of the individual nests in the large communal structures. As a matter of fact, the harmony of life within each colony, the lack of what may be likened to traffic congestion, i. e., the coming and going of birds in the task of providing food for the young, the fact that out of numbers of individual nests examined by various observers none were found with unusual numbers of eggs or young, all argue for individu- ality in nest occupancy. Whether each male has only one or several mates is, however, unknown. There have been several attempts to explain the structure of the large, composite nests of this species, some writers claiming that each pair of birds builds an individual nest, all of them close together, and then the flock builds the common roof over all the nests, while other writers have recorded that the flock builds a large structure and then each pair builds its individual nest into this structure. J never saw the actual beginning of a nest, and the smallest nests I found were, as mentioned above, complete structures with numbers of nests within them. However, Roberts (1940, p. 333) describes the construction of the communal nest as follows: . . . first a roof is constructed of coarse straws in the strong branches of a large... tree, and under this a great number of nest-chambers are made by nipping off the straws to form a tunnel upwards with a chamber at one side of the top of the tunnel; each pair of birds has its own nest-chamber, and scores of pairs may occupy the same colony. BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 305 To this I may add that regardless of how the first start is made, it is true that all through the nonbreeding season, the entire flock does a large amount of roofing and general enlarging of the whole affair so that it is true that subsequent individual nests are built into the large structure. The nests are added to year after year, and frequently become so large and heavy that they break the branches upon which they rest, and crash to the ground. All the birds seem to work together equally, apparently the males as well as the female(?)s, and even during the breeding season, when they have eggs or young in the nest, the male birds may be seen carrying straw to the roof or other parts of the common structure, not necessarily close to their own respective individual nests. The huge, massive affairs are composed wholly of dried grasses of a rather coarse, tough sort that grows commonly in southwestern Africa, and the seeds of which enter into the diet of the weavers very largely. The material is not really woven or even plaited on the surface of the nest, but is rather roughly put together in about the same way that hay is put into a well- made hay rack, but with a fairly definite thatching arrangement, causing the rain to run off and not soak through. The under side of the nest presents the rough, hard ends of the coarse straws and forms a very uneven surface. In the sparrow weavers and social weavers (subfamily Plocepas- serinae) we find, then, as far as our incomplete data permit us to generalize, an annual behavior cycle characterized by lack of migration or winter flock fragmentation, a substitution of a communal flock territory for individual ones as far as nesting is concerned, and a very marked development, both in seasonal duration and in individual activity, of the nest-building habit. The published observational data indicate that both sexes participate in nest building, but these data are open to question because of the similarity in plumage of the males and females; whether one or the other does most of the con- struction is not known. Nothing appears to be on record concerning the courtship habits, so it is not possible to ascertain whether this part of the cycle comes before or is associated with already completed nests as it is in some of the typical weavers (Ploceinae). Turning now to the next group, the weaver finches (subfamily Passerinae), which group includes the ubiquitous house sparrow (Passer domesticus) and its relatives, we find a different range of nest types. Some, like the house sparrow, build fairly bulky, formless, untidy nests in trees, on ledges, cornices of buildings, even in holes in trees, and other elevated sites (never on the ground). When built in the branches of a tree the nest usually is domed with an en- trance on one side, and fairly abundantly lined with feathers and other soft materials; when built in a hole the lining is much reduced as is 306 § ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 the rest of the nest structure. The birds are multiple-brooded; both sexes take part in nest building and in caring for the young. Although the nests are not such as would, in and of themselves, suggest that their makers were overly involved in building activities, there is evidence that in a closely related species, the Cape sparrow of South Africa, Passer melanurus, the birds use the nests throughout the year as sleeping places “ . . . especially in winter, when nests with more warm material are often specially built for the purpose” (Roberts, 1940, p. 334). In other words, in this species we find some indication of nest-building activity outside of the breeding season. Whether this is true for other forms of the genus is not known. To return to the house sparrow, the reproductive behavior cycle, as reported by Jourdain and Tucker (1938, pp. 157-158), is quite peculiar and is still in need of further study before it can be properly interpreted. The— . . . prominent feature of breeding-season is noisy display, in which sometimes one, but commonly several males hop with loud chirpings, round female with elevated bill and tail and drooping wings, but merely elicit pecks from irritated hen . . . Whole performance commonly ends with sudden dispersal of participants and appears unconnected with coition or even pairing. Gengler relates latter to rough-and-tumble scrimmages between several males without display, female commonly becoming involved as well, though selection of mates as result of these tussles seems not very clearly demonstrated. Coition is normally solicited by female with drooping wings and twittering note, without display by male, and may be repeated as many as a dozen or fifteen times in succession. Same observer states that both mated and unmated birds of both sexes are involved in displays, but that mated males display only to other females, never their own. He inter- prets display as relict of former genuine courtship, now functionless except as outlet for persistent display instinct . . . exceptionally coition may be preceded by typical display of male without usual solicitation of female. There is evidence, as well, that the species has a polyandrous or promis- cuous tendency, and Thompson (quoted by Jourdain and Tucker) considers the noisy displays are explained partly by this tendency, and partly by the males coming into breeding condition before the females. Other members of the subfamily, such as the yellow-throated spar- rows of the genus Petronia, the rock sparrows of the genus Gymnoris, and the gray-headed sparrow, Passer griseus, appear to nest chiefly if not wholly in holes in trees, in old woodpecker or barbet holes, or even in suitable natural holes of not too large a size. They generally line these nesting holes with fibers and feathers. The gray-headed sparrow has adapted itself to human habitations and frequently nests under the eaves of buildings. The chestnut sparrow, Sorella eminibey, not infrequently makes use of old nests of other weaverbirds although it does at other times build for itself. The absence of adequate data on the members of this group, other than the house sparrow, makes it impossible to generalize on any broad BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 307 scale or sure foundation. It is safer, then, merely to summarize the picture in the one relatively well-known member. The picture of courtship display is markedly altered from what we are in the habit of considering normal for most passerine birds—males displaying to any females but their own mates; females apparently soliciting rather than permitting coition, a precarious monogamy with a tendency toward polyandry and promiscuity. The scaly weavers of the genus Sporopipes form a subfamily by themselves, the Sporopipinae. They are not too well known, but I have found them in very loose flocks or small assemblages in the dry thornbush veldt of the Transvaal, where they feed on the ground like the Passerinae. The South African species (Sporopipes squami- frons) breeds during the southern winter as a rule, but at times during the summer as well, suggesting a not too well delimited nesting time. These birds are not colonial breeders, but build their roughly globular nests of grass stems and fine twigs, with a fairly pointed lateral en- trance, in the middle of the dense thorny branches of shrubs and low trees. Two nests that I found were less globular than published descriptions indicate is usual. They were somewhat similar to the untidy structures of the house sparrow, but smaller, slightly more compact, and less irregular in shape. In my field notes I described them as horizontal cylinders rather poorly closed at one end, and made of grasses, fine twigs, straws, etc. One nest containing three eggs was being very timidly guarded by two of the birds, presumably a pair (the sexes look alike). The birds would not stay near the nest while I was close to it, but returned to it as I walked away. Nothing seems to be on record concerning courtship, mating, or territorial behavior in any of the scaly weavers. We now come to the subfamily Viduinae, the indigo birds and the widow birds, containing a dozen species, three of which are definitely known to be parasitic and the others are suspected of having similar breeding habits. This group is somewhat intermediate between the Ploceinae and the next subfamily, the Estrildinae. Like the mem- bers of the Ploceinae, the Viduinae take 2 years to acquire adult plumage, and do not breed until then (the Estrildinae breed when 1 year old, as do the majority of small passerine birds). The adult males have a breeding plumage in which they are very different from the brown, streaky hens and year-old birds, the former of which they resemble in the nonbreeding plumage. (The Estrildinae do not show any seasonal plumage change as arule.) The best known of the Viduinae is the pin-tailed widow bird, Vidua macroura, and the fol- lowing description of its habits is taken largely from my field notes coupled with pertinent data in the literature. Vidua macroura is a gregarious bird and is usually seen in flocks of from 5 to 50 birds, depending on the season. In the breeding season 308 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 in South Africa, where seasons are definite, the flocks tend to break up and the birds pair off more or less. Yet it is not uncommon to see small flocks all through the breeding season. Such flocks usually contain but one full-plumaged male and the rest. of the birds are in the brown hen type of plumage. In some cases I shot into the flocks and found that the brown birds were year-old males, but in two cases the birds proved to be females with fairly enlarged ovaries. It seems, therefore, that this bird is somewhat polygamous, although I should judge from most of the cases I have observed (and they are many) that it is frequently, if not usually, more or less monogamous. In equatorial Africa all the individuals of the species in any one locality do not breed at the same time and these flocks usually contain a breeding pair and either year-old birds or nonbreeding adults. The lack of definite seasons complicates things superficially to the extent that the apparent state of affairs has no real relation to the actual conditions. This widow bird is largely terrestrial in habit and gathers most, if not all, of its food on or near the ground. However, in Natal, at least, during the southern winter the birds go about in large flocks and spend much time in the trees, where they act and sound not unlike small finches such as the North American redpoll, Acanthis linaria. They are by no means confined to trees and are found in tall grass and in reeds along stream banks. During the breeding season the males often use isolated trees as perches from which to sing and to watch over their territories, but the birds spend by far the greatest part of the time on the ground. On November 24, at Woodbush, Transvaal, I saw an adult male in full breeding plumage. It was perched on a bush in an open grassy field, and as I approached it flew off to a nearby bush and then to another not far off as I came close again. It made a small circling flight and came back to the original bush. On and off during the rest of the day I found it there each time I visited the spot and found by repeated trials that it could not be induced to leave it. It had definitely established its territory there, and apparently the bush in which it was first found was its singing perch. The next day I spent a couple of hours watching it and tried to make it fly off, but it would not go more than a hundred feet and then circle back gradually. There was a single hen bird in the immediate vicinity. I shot the male and found the testes were much enlarged. The plumage was still very fresh; in fact the long central rectrices still retained a little of their sheaths and one of them was so loose that it came out when I skinned the bird. In the same region I watched two other males that were established in their individual territories. One of them was watched for 3 successive days and was apparently without a mate as yet. It had a BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 309 territory about 400 yards in diameter, considerably larger than that of the first male, but more open, less bushy, and probably contained possibilities of no greater number of nests to parasitize than the other. The third and last male had a smaller breeding area and was usually accompanied by three or four brownish henlike birds. I shot one of these birds and found it to be a male—a year-old bird in first nuptial plumage. The courtship displayed was first observed at Woodbush, Transvaal, on December 1. The male flew up from the ground and hovered about 2 feet in the air directly over a female, with his body feathers shghtly ruffled and his wings beating rapidly. With each wing beat the four long rectrices were violently jerked and made to stream boisterously over the female, much after the cascade type of tail display of Coli- uspasser ardens and Coliuspasser procne. On another occasion, in equatorial East Africa, I saw a male display to a female that was perched in a thorn tree. The display was similar to the one already described; the male danced in a stationary posi- tion as though suspended in midair a couple of feet above the female. On still other occasions I watched males courting when there were several of the brownish hen-feathered birds present. In all such cases I noticed definitely that the male tended to confine his atten- tions to one particular bird. It seemed as though there was but one female and that the other brown birds were year-old males. In one case I shot the whole band (five brown birds) and found that one was a female in breeding condition and the rest were young males. Inasmuch as this widow bird is parasitic in its breeding habits it is interesting to compare it with the cowbirds (Méolothrus) of the Americas. The chief difference seems to be in their sexual relations. Both are more or less monogamous but the Vidua tends toward polygamy while the Molothrus tends toward polyandry. The vocal efforts of this species are not remarkable. The usual call notes are weak, high, but sharp tsips, something like the weaker notes of the redpoll (Acanthis linaria). When a band of birds calls simultaneously and rapidly they produce a light twittering chorus. The song is a rapid but modulated repetition of the call note and usually consists of from 5 to 10 syllables and occasionally more. It is given in flight as well as when at rest. Curiously enough, I never heard a male sing while going through his display dance before a female. As is well known, this species is parasitic in its breeding habits; i. e., it lays its eggs in the nests of other birds and leaves them to their care. Vidua macroura is not the only ploceid exhibiting this habit— V. regia and V. paradisea and, as we have already noted, Anoma- lospiza imberbis are also parasitic, and probably the other species of Vidua will in time be found to be parasitic as well. Vidua 310 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 macroura is parasitic chiefly on waxbills and generally lays but one egg ina nest. I have seen sets containing two, three, and even, four of the widow bird’s eggs along with those of the victims, but such sets are not usual. The eggs are pure white and differ from those of the common fosterers only in size. The following birds have been found to be parasitized by Vidua macroura: Lonchura scutatus Estrilda rhodopyga Estrilda astrild Estrilda delamerei Estrilda subflava Lagonosticta senegala Estrilda melpoda Lagonostica rubricata Estrilda massaica Amauresthes fringilloides Estrilda melanotis Coliuspasser ardens The incubation period is 12 days. The breeding season in South Africa is late in the southern summer— January, February, and early March, sometimes earlier. In Kenya Colony the species breeds during both the short and the long rainy seasons. The short rains come in November, December, and Jan- uary; the long rains in April, May, June, and July. As one goes northward the rains shift to later in the calendar year; thus in the southern Sudan the long rains extend into September and start correspondingly later than in Kenya Colony. The young V2dua does not always crowd out or starve out its nest mates (at least in the few cases I watched) as do the young cuckoos and cowbirds in so many cases, but all grow up together. Fully fledged young Vidua macroura are often found in flocks of young waxbills after leaving the nest but they do not remain long in these assemblages. Before they get ready to molt (postjuvenal molt) they form flocks of their own. I have seen as many as 15 or 20 young pin-tailed widow birds together. Frequently one or two adult birds, often males in breeding plumage, are found in these flocks. My observation on Vidua regia, the shaft-tailed widow bird, Vidua jischeri, the straw-tailed widow bird, and Vidua orientalis, one of the indigo finches, while much less complete than those on Vidua macroura, also indicate that the superficially apparent polygamy is actually not real, that while one male in adult breeding plumage may be accom- panied by a small flock of brown henlike individuals, most of the latter are immature birds of both sexes and only one in a group may be an adult female. In the case of the straw-tailed widow bird, Vidua fischeri, I once observed what seemed to be a territorial fight between two males in full breeding plumage. To summarize the behavior-pattern cycle in the Viduinae, we may characterize it as follows: apparently monogamous and solitary(?), but solitary only with respect to its own age group (adults), not solitary CLIGI ‘ZG “1OA ISI] “YBN ‘snyy ‘sowy *[INg ‘urdeyg wo17) SNWIMYADIN SNADO1d AO ANONOD ‘Z SNLVYIINOND SNADO1d AO ANONOD *} \ SH abe uuPWpallJ—"6p6| ‘ode UeluOsy WIG SHONVY LNA ISAN IVAGIAIGN| MOHS OL LSAN 4O ACIS YSGNN ‘Zz SNIDOS SNYIVLATIHd 3AO LSAN ‘1 ¢ ALV 1d uuewipatl.J—6p6| ‘Oday uPIUOsYyzIUIG LSAN TIWNOWWOD SIL -NVSIDS S3HL AO YANHOD V YSGNN SHAGTING AHL AO ANO “2 SNIDOS SNYIVLATIHd AO LSAN TIVWWS ATYIVSA GNV MAN ‘1 © saLvad uuewipalJ—¢6p6| ‘Woday weruosyytWIG Smithsonian Report, 1949.—Friedmann PLATE 4 1. A VERY LARGE OLD NEST OF PHILETAIRUS SOCIUS, PARTS OF WHICH HAD FALLEN DOWN BY THEIR OWN WEIGHT oe sss a 2. THE SOCIAL WEAVERBIRD, PHILETAIRUS SOCIUS Smithsonian Report, 1949.—Friedmann PLATE 5 foe — > j Pd 1. NEST OF PLOCEUS OCULARIUS (From Chapin, Bull. Amer. Mus. Nat. Hist., vol. 37, 1917.) 2. NEST OF EUPLECTES FLAMMICEPS (From Chapin, Bull. Amer. Mus. Nat. Hist., vol. 37, 1917.) Smithsonian Report, 1949.—Friedmann PLATE 6 Ry Upper, dancing ground of Coliuspasser jacksoni; middle, male Coliuspasser jacksoni on its dancing ground; lower, male Coliuspasser displaying to female on dancing ground. (All photographs on this plate from Van Someren, Journ. East Africa Nat. Hist. Soc., vol. 18, 1945.) Smithsonian Report, 1949.—Friedmann PLATE 7 1. CUCKOO FINCH, ANOMALOSPIZA IMBERBIS (From Shelley, Birds of Africa, vol. 4, 1905.) ’ 2. YOUNG ANOMALOSPIZA BEING FED BY PRINIA FALVICANS (From Roberts, Ann. Transvaal Mus., vol. 5, 1917.) VISSaY VNGIA AO ANO AGNV NMO Sl] 340 S999 SSaYHHL HLIM SNVSOIAV 14 VINIYd AO LSAN °Z SNLONOTMIdS SNADO1d AO ANONOD °| 8 31V1d uueulpaliJ—'6p6| ‘j40dayy ueluosyyIUIg BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN oll as far as immature “hangers-on” are concerned; courtship display well developed in all species; nest-building, incubation, and rearing instincts completely lacking in three members (V. macroura, paradisea, and regia) and probably in the others as well. The young of the three known parasitic species do not seem to evict or to starve out their nest mates of the host species, but may grow up in apparent amity with them. Roberts (1939, pp. 106-107) finds that while this is so, the female parasite usually destroys an egg of the host when depositing its own in the nest, but no such observations have been published. Usually there is but a single egg of the parasite in any one nest, but Roberts has found one instance where “‘five eggs of the common waxbill were all replaced by eggs of the Pin-tailed Widow-Bird.” Delacour and other writers have implied that the Viduinae are parasitic chiefly on waxbils, and even go so far as to suggest that each of the Viduinae has its particular Estrildinae host species, but this is by no means definitely established. Thus, the pin-tailed widow bird is known to parasitize at least nine species of Hstrilda and Lagonosticta, and two ploceine weavers, Coliuspasser ardens and Amauresthes fringilloides, while there is some evidence that Vidua regia lays its egg in the nest of a warbler, Prinia flavicans. The last subfamily of weaverbirds, comprising the waxbills, grass finches, and mannikins, is the Estrildinae. Delacour (1943, pp. 71-72) has recently summarized the characteristics of this group as follows: Small weaver-finches of highly specialized color pattern, never showing a primi- tive streaked sparrow-like brown plumage and horn-colored bill; sexes alike or different; immature always different from adult females. No eclipse plumage in males, with one exception. Nestlings always showing brightly colored, swollen spots, lobes or bands at the gape, and an ornamentation of the tongue or palate, consisting of spots or lines. Eggs numerous and always white; nests globular with a side entrance, but not woven. Young birds become adult within a year of their birth and are then able to breed, while it takes two years for young Viduinae and Ploceinae to mature. Peculiar song and courtship variable but consistent, in a general way, in large groups of genera. Ten primaries in the wing, the first being very short and falcate, with the exception of two genera (Clytospiza and Spermophaga) where it is moderately long, not parasitic. This large group is composed of three natural subdivisions: the wax- bills, chiefly found in Africa, but with one genus in Asia; the grass finches, found in Australia and some of the islands of the south Pacific; and the mannikins, found in Africa, Asia, and Australia. The Estrildinae never weave elaborate nests like the Ploceinae but con- struct roughly globular nests of grasses and leaves, with the entrance on one side, and which are usually built near or on the ground, in the grass, oF in bushes and low trees. The nests are very large for the size of their builders. A number of species frequently use old nests of other weavers, but usually do a certain amount of work on the 866591—50-——21 312 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 nests themselves, such as adding to or rearranging the lining. Thus, the bronze mannikin, Lonchura cucullatus, sometimes builds its own nest, but often breeds in old nests of other weavers, particularly of species of Ploceus. At Kaimosi, in western Kenya Colony, I found a rather untidy, loosely constructed nest of dried grasses and plant fibers, lined with grass seed-heads and feathers; it contained four white eggs and was evidently the nest of a pair of these mannikins, which were constantly seen on or about it. On the other hand, a day later I saw a Lonchura enter an old Ploceus nest, and, wondering what the bird might be doing there, I cut down the nest and found in it two eggs exactly like those found in the other nest the day before. The bird acted in a very excited manner as I examined the nest. Jackson (p. 1473) also records that this species breeds in old nests of Ploceus reichenowi, which it lines with grasses. Other Estrildinae known to use old nests of other species not infrequently are the silverbill, Lonchura cantans, the cut-throat finch, Amadina fasciata, the red- headed finch, Amadina erythrocephala, the common waxbill, Estrilda astrild, the zebra waxbill, Estrilda subflava, the lavender waxbill, Estrilda perreini, and the cordon-bleu, Uraeginthus bengalus. Aside from the fact that nest building in many of the Estrildinae is not so fixed in its pattern but that the birds may either build new nests for themselves or make use of old nests of other species (often very different in design from those their own species would construct), it is worth noting that in a good number of species the males take part in the task of incubating the eggs. Thus, in writing of the zebra waxbill, Jackson (p. 1517) goes so far as to say that “. . . as is gen- erally the case with Waxbills, the males assist in incubation.” Infor- mation on the courtship habits and sexual relations of the Estrildinae is still rather scanty, at least as far as significant and reliably worked-out details are concerned, but what data there are indicate nothing unusual in either respect. The birds appear to be monog- amous, and, as is so frequently the case with species in which the sexes look alike, the courtship antics do not show any peculiar or marked developments. In review, then, the great family of weaverbirds exhibits an aston- ishing range of diversity and variety in the mode of expression of the different parts of the reproductive behavior cycle. In the beginning of the breeding time we find everything from marked fragmentation of wintering flocks into pairs, to year-round gregariousness, and in courtship from a pattern that comes prior to nest building to one that follows the completion of the nests by the males, from solitary antics to elaborate display on special dancing grounds, and, on the other hand, to almost none at all, or, as in the case of the house sparrow, to barren but promiscuous displaying by mated males, seemingly BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 313 devoid of direct reproductive function, coupled with apparently monagamous coition-inviting display by mated females. Nest build- ing may vary from solitary to highly communal, and to none at all, and even to parasitism, from slovenly put together masses of material to amazingly fine and intricate weaving, or huge, communal super- structures, or may be reduced to merely relining a disused nest of another species or to lining a hole in a tree. Nest construction may be done entirely by the male, by both sexes, or largely by the female, or may be omitted entirely. Sexual relations vary from solitary monogamy or social monogamy to polygamy, polyandry, and to ap- parent promiscuity. Incubation in some species, or groups of species, is performed solely by the hens, while in others the cocks share the task with their mates, or, in the case of still others, neither sex takes any care of the eggs, but are parasitic. 'The members of the subfam- ilies Ploceinae and Viduinae do not come into breeding condition or acquire adult breeding plumage until they are 2 years old; the mem- bers of the other groups breed when 1 year old; this in itself is a pro- found difference. In some forms of the Viduinae and Ploceinae it permits a type of breeding-season gregariousness, although only a single adult male and female are usually involved in each little flock. Few, if any, families of birds offer such a bewildering array of variations of the parts of the annual cycle, and I cannot help but wonder if some of these variations may not have been due originally to the extremes to which, in previously established variations, some of the stages had been carried. At least the situations created by some of these extreme developments seem to have been propitious for further and even quite contrary subsequent changes. Paradoxical as it sounds, it is possible that the excessive develop- ment of the nest-building habit may actually have been a contributing factor in the origin of the complete absence of nest building and egg care that we know as brood parasitism. In cases of extreme indul- gence in nest construction such as we find in the social weaverbird (Philetairus socius) and some of its relatives (Plocepasser etc.), the huge bulky structures are added to, chiefly by the males, all through the nonbreeding season. By the time the birds are ready to make their own individual nest tunnels in the already existing superstruc- ture they are not acting very differently from birds that make use of old nests of other species which they then repair. In the case of the numerous species of typical weavers (Ploceinae) in which the not yet breeding males construct many nests, the subsequently mated females are again in a not dissimilar position of taking over nests which they themselves have not built, and relining them and breeding in them. It seems that Ali (1931) must have had some such thought in mind when he noted that Baya weavers occasionally laid eggs in disused 314 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 nests of others of their own kind instead of making new ones for themselves, as this prompted him to raise the following argument: If the bird laid in disused nests it would only succeed in avoiding the labour of building, but would still have to incubate the eggs itself. If on the other hand it was successful in slipping into an unguarded Baya nest whence the brooding hen had gone (as actually happened on September 18) and in laying its eggs there, it would be, quite involuntarily, but with good effect all the same, compelled to retire on the return of the legitimate occupant, leaving its egg to be hatched by the Baya. Would such a process not tend, in course of time, to develope into, and establish, a habit of systematic and voluntary parasitism as has been observed in some African weavers? In this connection it may be recalled that Lynes (1924, p. 661) found that in nesting colonies of several species of African weavers related to the Baya, studied by him in the Darfur Province of the Sudan, many nests contained one or two extra eggs of the same species as the host, but recognizably distinct by virtue of different color or state of incubation, in other words, eggs that probably were laid by other individuals of the same kind. It seems then, both in Asia and in Africa, that not infrequently female weavers, ordinarily using nests they have not built themselves, may lay an occasional egg in a nearby nest of their own species. The Viduinae are, as stated earlier in this paper, intermediate be- tween the typical weavers (Ploceinae) and the waxbills (Hstrildinae). In many species (perhaps the majority) of the former group, and also in a good number of forms of the latter group, the hens breed in nests, the actual construction of which has been foreign to their experience and their efforts; in many forms of the latter group, and at least some members of the former subfamily, the care of the eggs is taken over, at least in part, by the cocks. The parasitic mode of reproduction occurs, as far as known, in five widely separated and quite unrelated families of birds—the ducks, the cuckoos, the honey-guides, the weaverbirds, and the hang-nests (cow- birds). There can be little doubt that the development of brood parasitism has taken place independently in each of these five groups, and it is not without significance, or at least suggestive value, that this highly aberrant reproduction pattern has developed among the small passerine birds (generally considered to be the most highly evolved of all the birds) in those two families some of whose members have carried the habit of nest building to its highest and most complex development. It is all the more noteworthy that in the weaverbirds, a larger group than the hang-nests and one with greater diversity of behavior patterns, the parasitic habit has developed in two sub- families, apparently {independently—the cuckoo finch, Anomalospiza imberbis, in the Ploceinae, and in the members of the Viduinae, three of which are definitely known to be parasitic, and the rest of which BREEDING HABITS OF WEAVERBIRDS—FRIEDMANN 315 are strongly suspected of having the same habit. Many more details have still to be learned of the annual cycle of behavior patterns in these birds before it may be possible to attempt to determine the precise causes and the subsequent evolutionary paths that twice in the history of the weaverbirds have lead from nesting and incubation and caring for the young to a state of brood parasitism. REFERENCES Aut, Sauim A. 1931. The nesting habits of the Baya (Ploceus philippinus). Journ. Bombay Nat. Hist. Soc., vol. 34, No. 4, pp. 947-964. Bates, Grorce L. 1930. Handbook of the birds of West Africa. BELCHER, CHARLES F, 1930. The birds of Nyasaland. CuaPiNn, JAMES P. 1917. The classification of the weaver birds. Bull. Amer. Mus. Nat. Hist., vol. 37, art. 9, pp. 243-280. DELACOUR, JEAN. 1943. A revision of the subfamily Estrildinae of the family Ploceidae, Zoologica, vol. 28, pt. 2, pp. 69-86. Deacour, JEAN, and Epmunp-Buanc, F. 1933-1934. Monographie des veuves. Oiseau, n. s., vol. 3, pp. 687-726; vol. 4, pp. 52-110. FRIEDMANN, HERBERT. 1922. The weaving of the red-billed weaver bird in captivity. Zoologica, vol. 11, No. 16, pp. 355-372. 1929. The cowbirds. A study in the biology of social parasitism. 1930. The sociable weaver bird of South Africa. Nat. Hist., vol. 30, No. 2, pp. 205-212. 1935. Bird societies. Jn A Handbook of Social Psychology, chap. 5 (edited by Carl Murchison). JACKSON, FREDERICK J. 1938. The birds of Kenya Colony and the Uganda Protectorate. 3 vols. Jourpain, F. C. R., and Tucknr, B. W. 1938. In Witherby, Jourdain, Ticehurst, and Tucker, Handbook of British Birds, vol. 1, pp. 157-158. Lack, Davin. 1935. Territory and polygamy in a bishop-bird, Euplectes hordacea hordacea (Linn.). Ibis, ser. 18, vol. 5, pp. 817-836. Lynes, HuBERT., 1924. The birds of north and central Darfur. Ibis, ser. 11, vol. 6, pp. 661- 678. Prisst, Cecit D. 1933-36. The birds of Southern Rhodesia. 4 vols. Roserts, AUSTIN. 1917. Parasitism amongst finches. Ann. Transvaal Mus., vol. 5, No. 4, pp. 259-262. 1939. Notes on the eggs of parasitic birds in South Africa. Ostrich, vol. 10, pp. 1-20, 100-117. 1940. The birds of South Africa. 316 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 SHELLEY, G. E. 1905. The birds of Africa, vol. 4, pt. 1, pl. 31 (opposite p. 108). Srark, ARTHUR C. 1900. The birds of South Africa, vol. 1. TAYLOR, J. SNEYD. 1946. Notes on the masked weaver. Ostrich, vol. 17, No. 3, pp. 145-155, Van Someren, V. D. 1945. The dancing display and courtship of Jackson’s whydah. Journ. East Africa Nat. Hist. Soc., vol. 18, Nos. 3-4, pp. 1381-141. Van SomEREN, V. G. L. 1916. List of birds collected in Uganda and British East Africa, with notes on their nesting and other habits. Ibis, ser. 10, vol. 4, pp. 373-472. NEW ZEALAND, A BOTANIST’S PARADISE By Eaprert H. WALKER Associate Curator, Department of Botany, U. S. National Museum [With 10 plates] INTRODUCTION The wisest traveler learns as much as possible before a trip, sees all he can during his journey, and corrects and enlarges his knowledge by further reading and inquiry after returning home. This article is the outgrowth of the author’s short but full visit in New Zealand, which was ideal in nearly every respect except for lack of advance knowledge of the country, especially of its botany. It suggests what the writer would have liked to know in advance but had to learn on the trip and after it. The suggestions given here for further reading may be of interest, not only to the fortunate few who will visit New Zealand in person, but the greater number who may do so vicariously by reading and by listening to those who have gone. The plants of New Zealand can hardly fail to gain the attention of the visitor, and the student of New Zealand will find abundant refer- ence to them in his reading. People in an industrial country may ignore the plant life, but those in an agricultural land like New Zealand cannot escape the imprint of the vegetation on their lives. New Zealand is a land of enthusiastic and competent amateur naturalists, and its professional botanists are outstanding. The visitor will find a local naturalist in nearly every town or center, who is eager to share his specialties with the stranger and to show him the offerings of the field. ‘The traveler to New Zealand will probably first meet the introduced flora which dominates the landscape in the inhabited parts. Only when he visits the more remote and undisturbed areas will he see many of the native plants. If he has an economic or agricultural bias, probably the grasslands, the backbone of New Zealand’s economy, will impress him most. (See pl. 9.) If he is conservation-minded, the sight of the vast area of shrubland and fernland (Pterrdium aquilinum var. esculentum) will make him pain- fully aware of man’s destruction of the native forest. But the soul of the pure botanist, undisturbed by problems of economics and con- 317 318 | ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 servation, will be stirred most as he enters the forest or “bush” where abounds the native New Zealand flora, so rich in plants found in no other country—that is, the endemic species. (See pls. 2 and 3.) Besides the grassland, scrub, and bush he will see other plant forma- tions. The extensive plantations of trees, all planted in rows of uni- form age, are very impressive. The species so grown are all exotics, that is, not native to New Zealand, the principal one being the Monte- rey pine (Pinus radiata), a useless tree in its native California, but here by far the most economically important tree to be found. Not only does it occur in plantations, but it is to be found almost every- where as a hedge tree or windbreak (pl. 9, fig. 1) and even as a naturally planted weed invading wasteland. It is essential in understanding the peculiarities of the flora of New Zealand to know its location and climate as contrasted with that of more familiar areas. The vegetation or the major plant formations will then be discussed, after which the flora or the elements which compose the vegetation will be taken up. Finally some consideration will be given to the past and present study of botany in the country. LOCATION AND CLIMATE New Zealand extends from about the 34th to the 47th parallel south latitude, a distance of about 900 miles. (Fig. 1, and fig. 2, p. 333.) It consists essentially of three islands, North, South, and Stewart Islands, with a few small nearby islands or islets. There are several outlying island groups, politically and biologically part of New Zealand, of much interest, but they are not included in this discussion. For vivid geographical comparison, suppose the three main islands of New Zealand were inverted and superimposed on North America at a corresponding latitude, with the North Cape of North Island at Cape Lookout about the center of the Atlantic coast of North Carolina. Then the South Cape on Stewart Island would lie north of Quebec in Canada. The East Cape at the end of the Grisborne Peninsula of North Island would be in southern West Virginia, and Mount Egmont at Harpers Ferry, W. Va. Wellington, at the southern end of North Island, would be in west-central Pennsylvania, and Christchurch on the east side of South Island would be on the shore at the east end of Lake Ontario. Both areas are in temperate zones of the earth’s surface, but their climates are in striking contrast. Eastern North America has a continental climate with extremes of temperature and a moderate, irregular precipitation. New Zealand, however, has a strong oceanic climate with far milder temperatures throughout and a much smaller difference in temperature between the northern and southern ends and between winter and summer. The precipitation is fairly evenly distributed throughout the year, although it varies from place to place. New Zealand owes its climate to the unifying NEW ZEALAND—EGBERT H. WALKER 319 LS SO arr — 3 <5 --t-Fe- apricorrg "| 7 b; Db - a “7 N\ \ SS > =Chatharn & aN Na SER S|! Tee | SS 1Bocirity ves IES — Anti vOWeSN| eee aN °° Acwkleind [5.22 : wot a ace ! 4 | sas PH eg bos y —- 7GILUG 2 oF) A | Du lacguarrie ss ! Se eee | eZrneraldl [4 Ficure 1—Chart of Tasman Sea showing deflection of the cold Antarctic drift by the warm East Australian current from the Tropics. (From W. C. Davies (12).) influence of the warm Tasman Sea on the prevailing westerly winds which blow over it from Australia, over a thousand miles away. In its oceanic climate lies the explanation of many of its vegetational contrasts with other countries. The vegetation in northern New Zealand is far more tropical in appearance than its geographical counterpart in North Carolina. 320 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 This ‘subtropical’ aspect of the forests can be seen even in northern South Island and grades into the characteristic features of the dense temperate rain forests of the west coast of South Island. The south- ern-beech forests of Nothofagus in the south are clearly Temperate Zone forests. (See pl. 1, fig. 2.) The mountains in New Zealand cause more changes in the climate from place to place than does the latitude. The mountain ranges and plateaus of North Island le mostly east of the center. They are largely volcanic and influence the vegetation not only through their effect on the winds and moisture but also on the soil. The volcanic ash and pumice readily absorb more moisture, much of which seeps away beyond the reach of the plants growing on the surface. Mount Egmont is a majestic isolated volcanic cone on the west coast with vegetation in characteristic altitudinal zones from sea level to the perpetually snow-covered summit. In South Island the rugged Southern Alps parallel the west coast and thrust their peaks far into the zone of permanent snow. (See pl. 7, fig. 1.) Their highest peak is Mount Cook, its summit 12,349 feet above the Tasman seashore less than 24 miles to the west. They are formed by erosion of uplifted land rather than volcanic activity and are composed largely of friable greywacke rock. (See pl. 7, fig. 2.) These ranges drain the prevailing westerly winds of most of their moisture. Thus the west coast has a heavy rainfall of around 200 inches a year, while on the plains of the eastern leeward side there may be as little as 20 inches. The highest annual rainfall yet re- corded is 228 inches at Puysegur Point on the west coast, and the lowest, 13 inches, in Central Otago only 150 miles away. Dense rain forests cover the steep western slopes and the narrow coastal plain below, whereas on the east are the natural grasslands and the broad cultivated plains. In Central Otago is a semidesert area. The transition fromheavy rain and dense forest to sunshine and almost barren eroding slopes may be made in a surprisingly short time in driving over the divide. Other ranges and hills, especially the Kaikora Ranges in Marlboro and those in the rugged Banks Peninsula, inter- rupt this picture and diversify the ecological conditions and vegeta- tion. Thus New Zealand has a great variety of distinctive, plant formations in a remarkably small space, a fact which makes bota- nizing a most interesting and relatively easy occupation. The climate of New Zealand has been presented by Kidson (16, 17).! THE GRASSLANDS The grasslands of New Zealand are the foundation of its agricul- tural economy, and one is sure to be impressed by their extent and 1 Numbers in parentheses refer to the bibliography. NEW ZEALAND—EGBERT H. WALKER 321 . variation. They are of two kinds: first, the original tussock grass- lands of native species (pl. 9, fig. 2), and second, the pastures with a sward formed of introduced species (pl. 9, fig. 1). The history of New Zealand is largely the story of man’s replacement of the native bush with pastures and the exploitation of the native grasslands in feeding his flocks of sheep. In the ashes of the bush he planted grasses and began the process of adapting the sward-forming techniques, so well developed in his native England, to the conditions of this new and promising land. He fought a continual battle with the few native plants with weedlike tendencies and the more numerous and generally more vigorous exotics, which he intentionally or unintentionally brought from the far parts of the earth. From early blunderings he has now developed the technique of sward growing to a very advanced degree, and agricultural progress or deterioration in large parts of North Island and certain regions of South Island depends on the com- position and condition of this pasture sward. Much land formerly covered with fern, scrub, or blackberry is now grass-covered, with a high sheep-carrying capacity. It was a most enlightening experience to see the work of the Animal Research Station at Ruakura near Hamilton in Auckland Province, in breeding and mixing strains of grasses, on which, with proper rotation, an amazing number of sheep can graze throughout the year without additional feed. The various types of these artificial grasslands of North Island have been carefully mapped and analyzed in a publication by Madden (19). Something of the history and significance of these grasslands can be gleaned from the account written by an English‘agronomist, Stapledon, who visited Australia and New Zealand in 1926 (25). The planted pastures of South Island, developed by essentially the same means, are well described in Hilgendorf’s ecological survey of the grasslands (13), which supplements Madden’s. The natural grasslands are very different from the man-made pastures. They occur most extensively in South Island, but smaller areas are to be found in North Island, especially in the central plateau. A relatively low rainfall with a cooler and more even temperature are among the principal factors governing the development of grasslands rather than shrublands or “bush.’”’ The plant composition of this formation varies considerably according to local conditions. Pas- toralists generally recognize five tussock grasses. ‘Two of them, snow grass (Danthonia raoulii var. flavescens) ? and red tussock (D. raoulu var. rubra) are the tall tussocks, 3 to 6 feet high, while the short tussocks, 1 to 2 feet high, are the silver tussock (Poa caespitosa) and the hard or fescue tussock (Festuca novae-zealandiae). The blue tussock (Poa colensoi) is only 6 to 9 inches high. 2 The names, both common and scientific, in general use in discussing these grasses vary considerably. This makes research on the grasslands rather difficult. Zotov (28) recognizes only four tussock grasses. 322 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The history of the tussock lands is an almost continuous story of progressive deterioration due to overgrazing, burning, rabbit infesta- tion, and increased wind erosion, land slip, soil creep, and water erosion. The present deserts in northern Canterbury, Marlboro, and Central Otago Provinces were grasslands when white settlement began, and the carrying capacity of most sheep runs is today far less than it was in the beginning. Everywhere one can see erosion that is of recent origin. A most definite sign of overgrazing in South Island is the excessive development of the scabweed, Raoulia lutescens. (See pl. 8, fig. 2.) Usually the rabbit population increases as the tussocks diminish and more open spaces are formed. This only accelerates the destructive process. It is important to keep in mind that the tussock grasses themselves are rarely grazed, except the new growth which springs up after they are burned over, which is tradi- tionally done annually. The role of the tussocks is to furnish pro- tection to the smaller grasses and other plants which grow among them and furnish most of the feed. The tussock grasslands of South Island have been dealt with rather fully by Zotov (28) and recommendations presented for restoring these areas to production. First, annual burning must be eliminated or, if absolutely necessary to eliminate shrubby invaders, replaced with carefully controlled burning. The number of grazing animals must be reduced to the carrying capacity of the land and rotational grazing introduced in order to restore the fertility. When necessary, the tus- sock grass must be replanted with selected unpalatable strains or jor- danons, and, when a protective covering is thus established, highly palatable strains of native species must be sown between the tussocks. Hardly any of these measures are now used by the sheepmen. It was most gratifying to have a glimpse of the Government’s research work in tussock-grass restoration at its field station in Hutt Valley near Wellington. The work done there is preliminary to research and experimentation in the tussock country itself and will surely some day result in restoration of much depleted land. Probably some areas have gone almost beyond reclamation and will remain, as have so many other parts of the world, monuments to man’s lack of fore- sight and self-control in seizing all the produce of the land rather than just its surplus. SHRUBLAND AND FERNLAND No traveler in New Zealand can fail to be impressed, and at the same time generally depressed, by the vast extent of land covered by shrubs and ferns. Unlike the grasslands, they bring no sense of well- being to man, and, compared with the forest, they at first seem botanically unattractive. But neither impression is wholly correct. NEW ZEALAND—EGBERT H. WALKER 323 Shrubland is any plant community in which tall trees are wanting and shrubs dominate. The fern, which comprises the fernland, is the native variant of the world-wide bracken, Pteridium aquilinum var. esculentum, or P. esculentum of many authors. Fernland is here linked with shrubland because the fern reaches shrub size and the formation is as dense and impenetrable as the densest thicket of woody shrubs. Furthermore, this fern community is closely related ecologically to the other most extensive shrub community, that dominated by manuka? (Leptospermum scoparium or L. ericoides— Myrtaceae) (pl. 5, fig. 2). Together these two cover more area than do the other shrub formations, of which there are many in very different habitats and of diverse composition, form, and origin. As in other lands, these shrub formations develop in response to certain natural conditions. These conditions may develop over a long period of time. When sudden changes occur favorable to the growth of a shrub community the formation is called an induced formation. Shrub formations follow certain volcanic eruptions and sometimes floods or places of excessive erosion. But more significantly they are man-induced, coming along after the forest has been destroyed with ax and fire, and grass has been sown on the ashes, or where man’s fires and his overabundant greedy sheep have destroyed the natural grass cover. (See pl. 4, fig. 1.) The manuka (Leptospermum scoparium—Myrtaceae) is a shrub or small tree with an amazing adaptability and persistence. It usually forms a community without the bracken or it may be variously mixed with this fern. It seems able to grow anywhere, wet or dry, in good soil or bad, and in heat or reasonable cold, but not in alpine con- ditions or deep forest shade. Its outstanding ability to thrive on poor soil makes it rush in where man has done his best to destroy the land. It is especially prominent on the gumlands of North Auckland, dug over and the fertility dissipated in the search for fossil kauri gum, desired as an ingredient in high-grade varnish. Manuka is extremely plastic in its response to its environment. Within this community there are some 81 other species of plants, many of them of great interest to the botanist, not the least being the bulbous-rooted New Zealand orchids. Manuka is an important source of fuel for man, 3 To the foreigner the widespread use in New Zealand of native Maori names for trees and other plants is somewhat disconcerting. Very often there is no other name, as for example, manuka for Leptospermum scoparium or kauri for Agathis australis. These names have often been made into the specific scientific names, as taraire in Beilschmeidia taraire and tawa in B. tawa. Another common practice of many New Zealanders is to use in speech a specific scientific name for a common name, as macrocarpa for Cupressus macrocarpa, lawsoniana for Cupressus lawsoniana, and radiata for Pinus radiata, the Monterey pine. Another disconcerting practice is the use of some common English name, such as pine, for something which a foreigner at least would hardly recognize as such. Native pine in New Zealand refers to species of Podo- carpus. ‘Thus the traveler is in a new nomenclatorial atmosphere. For popular names of New Zealand plants see Andersen (2) and Cheeseman (4). 324 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 and in its shade grow seedlings of many forest trees, which eventually rise up and wipe out this ‘‘nurse” species by overshading it. So the manuka shrubland fills a varied and not altogether harmful place in New Zealand’s plant economy. One cannot feel quite so resigned to the fernland, though one quickly does resign from the job of trying to penetrate it. Its adapta- bility and prolificness closely matches the manuka, but within its dense growth there are few if any other plants. Like the manuka, it cannot endure much shade or cold, and so is not a denizen of the forest and alpine or subalpine slopes. Man may burn its tangled fronds, but new ones rise quickly from the unharmed underground stems. However, overstocking with cattle which eat the tender young fronds catches the bracken in its “tendon of Achilles,” and if the practice is persisted in, this scourge can eventually be conquered. A third shrubland community of much prominence is not only the result of man-made conditions, but of man’s introduction of plants. The English gorse, Ulex europaeus, was first brought, no doubt, to relieve man’s nostalgic longing for the lovely English countryside and to lend color to the generally colorless New Zealand vegetation. But this was an imprudent act. From the hedgerows it spread easily to adjacent fields, dry, gravelly river beds, formerly forested hillsides, and pastures. It is quite indifferent to the quality of the soil. Large open spaces soon became impenetrable thickets. Man constantly burns it off, but fire seems only to improve the viability of its seeds and to impoverish the soil, which harms the accompanying fodder plants more than it does the gorse. Control over large areas by grubbing it out of the ground is hopeless in this land of limited labor and large demands on human resources. Handcuffed with this gorgeous yellow culprit are the broom (Cystisus scoparius—Leguminosae), the rose, and the blackberry, and several adventive shrubs from adjacent Australia, especially hakea (Hakea acicularis—Proteaceae). One American shrub of this category is the tree-lupine of California, Lupinus arboreus, brought as a sand binder and now spreading beyond its first plantings to other sandy and gravelly spots, not always according to man’s wishes. The term “scrub” is often applied to any shrub formation. In Australia it is erroneously applied to certain forest formations, but Cockayne (7) applies the term in a more restrictive sense to any community of divaricating, stiff, shaggy, and often spiny shrubs. Such shrubs have numerous extremely wiry or rigid, much interlaced branches and twigs which zigzag at a wide angle in every direction. The thickets they form are close, unyielding, and often cushionlike masses. ‘To push ones way through this scrub is impossible and to travel over it is often a hazardous undertaking. The scrub is usually subalpine and is composed of various species of Coprosma (Rubiaceae), NEW ZEALAND—EGBERT H, WALKER 325 Cassinia and tree-daisy (Olearia—Compositae), wild Irishman (Dis- caria toumatou—Rhamnaceae—pl. 6, fig. 1), and Myrtus (Myrtaceae), though altogether there are about 55 species in 18 families which have this divaricating habit. The plant collector who makes pressed specimens almost meets his Waterloo when he tries to make a herbarium specimen to represent adequately such a plant. Some divaricating shrubs further thwart the collector by dropping their leaves, flowers, and fruits almost at the first gentle touch. The divaricating branches of pohuehue (Muehlenbeckia astoni—Polygonaceae) are pliant enough, but when a representative specimen has been warped into a plant press there is rarely a leaf or fruit left in situ, and a vivid supplementary description is needed to bring to the observer’s mind any adequate concept of the original habit of the plant. One will find various shrub formations in a wide range of habitats. Besides the extensive hillside formations of manuka and fern, and the subalpine scrub, this type of vegetation is often found on sea coasts (both rocky and sandy), wet lands, mineral lands, areas of volcanic ash and pumice, and wind-swept shores and mountain slopes. Its component species, growth forms, and adaptations to environmental conditions are of much interest. In some places associations of trees, dwarfed to shrub size by wind, salt spray, or soil influences, resemble and merge into shrub formations. FORESTS The principal natural resource in New Zealand when the pakeha or white man first came was its trees; at the present time it is its grass. But the white man could live only secondarily on the forest, so the trees had to go in order that he might provide for his primary need— food. Hence, this natural resource, which formerly covered almost the whole of North Island and much of South Island, was sacrificed at a rate hardly equaled anywhere else in the world. It took Europe four centuries to exploit its forests and America two centuries, but New Zealand accomplished this in one century. Although the exten- sive natural forests of the past are gone, the remaining fragments are sufficient to tell us a great deal about New Zealand’s botanical history. According to Cockayne (7), 385 species of plants are characteristic of the forests. Of these, 99 are trees, 63 shrubs, 51 herbs, 26 grasslike plants, 88 ferns, 26 climbing plants, 15 epiphytes, and 13 parasites. Ninety percent are endemics. So it is in the New Zealand forest that a visiting botanist will find his greatest delight in becoming acquainted with the true New Zealand flora. If he arrives in North Island, as is most likely (and much regretted by the people of South Island), his first acquaintance will almost surely be with the rain forest, which is composed of many species of trees and shrubs of many genera and 326 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 families. (See pl. 3, fig. 1.) Later he will meet the strikingly dif- ferent southern-beech forest, composed almost entirely of one or two species of Nothofagus and few shrubs. (See pl. 1, fig. 2.) There, if he is from the Temperate Zone of North America or Europe, the traveler will feel much more at home, although all the plants will be new to him. The rain forest in New Zealand is clearly tropical in its origin and affinities; indeed, it is often designated as subtropical rain forest, although it lies entirely in the Temperate Zone. Its character is the result of the oceanic climate with its mild, rather uniform temperatures, and its abundant, evenly distributed rainfall, which assures a high atmospheric humidity throughout the year. The southern-beech forest of Nothofagus, however, is clearly temperate in origin, with its nearest affinity the Nothofagus forests in Patagonia, Tierra del Fuego, and southern Chile, on one side, and in Tasmania and adjacent Australia, on the other. This forest occurs in New Zealand in cooler and less humid regions than most of the rain forests. It depends more on ground water and thrives in a less humid atmosphere. All the forests in New Zealand are evergreen. There are a few deciduous trees but none of them are forest dominants, so there are no deciduous forests in New Zealand. The rain forest is complex, the beech forest relatively simple. These two forests may be contrasted in part, as follows: Rain forest Southern-beech forest Tropical in appearance. Temperate in appearance. Composed of many tree species in many Composed of one or two tree species in genera. one genus. Dense within. Open within. With several plant strata.‘ With only an open layer of shrubs be- tween canopy and ground. Bases of trunks often with plank but- Trunks without buttresses. tresses. With many vines or lianas. With very few vines. Loaded with epiphytes. With only a few parasites. With many ferns. With few ferns. Each forest formation has various forest associations within it, these being quite complex in the rain forests and relatively simple in the southern-beech forest. The forest associations are named according to the dominant species within them, the principal associations in the rain forests being: (1) kauri (Agathis australis—Pinaceae), (2) mixed dicotylous-taxad,®? and (3) “white pine” or kahikatea (Podocarpus dacrydiovdes). 4 “Forest is piled upon forest.””—Humboldt. 5 Cockayne’s term for this forest association varies. It is called a mixed taxad forest (6) and dicotylous- podocarp forest (7). Species of New Zealand “pine,’’ Podocarpus or Dacrydium, members of the yew family Taxaceae, are dominant or characteristic. There are also many trees of various families of dicotyledons, whose seeds have two cotyledons in contrast with the conifers which have several and the monocotyledons with only one cotyledon. The use of the term dicotylous rather than dicotyledonous is in conformity with Cockayne’s usage (7). Smithsonian Report, 1949.—Egbert H. Walker PLATE 1 ] 1. LAKE FERGUS NEAR THE HOMER TUNNEL, WESTERN OTAGO, SOUTH ISLAND Southern-beech (Nothofagus) forests clothe the mountains high up toward the snowy peaks of the Southern Alps. 2. SOUTHERN-BEECH FOREST (NOTHOFAGUS FUSCA), PARADISE, LAKE WAKATIPU, OTAGO PROVINCE, SOUTH ISLAND Without looking at the foliage one would think himself to be in a northern-beech (Fagus grandifolia) forest in temperate North America. Cseraed O° M Aq ydeiz0j04q) ‘a[sunf[ esuap siqq 9z119}0BIBY CS|R soyAydide juespunge pue (jJa[) snuepUed SUIGUII[D 9q,J, *901} ISaI0} (‘UOSTON ‘AIN{YSUT U9IQYMBD Asoeqin>+o ‘seraeqd “OM AQ esny 8 8q 0] SMOIS puB “4SOY S}I Se[suRIys pus SdojaAus pue ‘punois qdeis0j0yd) 480} LOT UM0ID 94 JO pwaids 9Y} PUR J9eJ G9T SI IYSIaY [eI0} 94} 0} UMOP sayover ‘ayAydide ue sv ast SuIseq Al[[ensn BIBI BYAL aL ‘SeyouBsq IsIY ay} 0} YSiY JoaJ QO PUB JOJBUIBIP UL Joa] 44 fT SL 80 Uy, LSSYO4 AVXVL-SNOTALOSIC GNV1IS| HLYON ‘GNVY1TIHONY HLYON ‘LSSYHOS GaXIW SHLNI (VLSNEOY SONSCISOYLAW) VLVY V “Zz S3LVLSVNOdIVM AHLNISSYLIYNVy Vv... ALNHYWANVL,, ‘1 P SINAN J9¥]2A\ H 42934 —6h6| “HOdey weruosyztUIg ae ¢ 3LV1d Cprey sf 4q ydeisojoyg) * (44811) HOOT [BIIOA B SULIAAOD (Digns DYNnODY) Caeys as[qejeseA Useds pues -(punoise10} pue Jeyued) “ds pisniydng ‘(4Jap) s271qvzoeds visiw7ag GNV1S| HLYON “AONIAOYd NOLONITISM ‘SONVY WOVE _L SINV1d HOOY NIVLNNOW SAO ASTGAW V °Z € ALV1d (sarang "DO °M Aq ydeis -0JOUq) *4S910j YOIISIYI JO BIBIYS [RIBAS BY BSIIdUICD SIIqBY asIaATp JO se1oeds gel] Auvu pue ‘soyAydide ‘seuely ‘sued neyiu 04} ‘SLLI8} BOL, 1SSHYO4 AVXVL “SNOTALODSIG GSXIW AHL NI HLMOYUDS SSN3Q AHL ‘ff JONIeM “LI 1100G39—*4b¢4)] ‘qu0dayy UPIUOsSU]IWIC Smithsonian Report, 1949.—Egbert H. Walker PLATE 4 1. EDGE OF WAIPOUA STATE FOREST, NORTH AUCKLAND, NORTH ISLAND The untouched primeval kauri forest (right) formerly extended over the now cut-over stump, bracken, grass, and exotic shrub covered sheep-grazing land (left). (Photograph by E. H. Walker.) 2. COASTAL DYSOXYLUM SPECTABILE FOREST, STEPHENS ISLAND, COOK STRAIT The forest floor is free of undergrowth, probably partially damaged by cattle. The Dysorylum has characteristically twisted trunks and above-ground roots. The slender trunks are Piper eacelswm. (Photograph by L. Cockayne, courtesy New York Botanical Garden.) Smithsonian Report, 1949.—Egbert H. Walker PLATE 5 1. TREE FERN, SHORE OF LAKE ROTO-ITI, NORTH ISLAND Large tree ferns of many kinds are denizens of the forests and cut-over lands, and often of the roadsides i je toe Bye ae ees os 0 eer i 7 Pe 2. MANUKA (LEPTOSPERMUM SCOPARIUM), EAST COAST SOUTH ISLAND This abundant association clothes vast open lands from sea coast and swamps to mountain sides with an almost impenetrable thicket. (Photograph by L. Cockayne, courtesy New York Botanical Garden.) Smithsonian Report, 1949.—Egbert H. Walker PLATE 6 1. WILD IRISHMAN (DISCARIA TOUMATOU), A DIVARICATING SHRUB, DRY EAST COAST OF SOUTH ISLAND This characteristic growth habit is found in many New Zealand shrubs. See also plate 7, figure 1. (Photograph by L. Cockayne, courtesy New York Botanical Garden.) 2. COLLECTING DONATIA NOVAE-ZELANDIAE IN AN ALPINE BOG, MAUNGATUA RANGE, OTAGO PROVINCE, SOUTH ISLAND This dense flat mat is so solid that footprints hardly show. Other species of this genus occur only in Tasmania and southern South America. (Photograph by E. H. Walker.) Smithsonian Report, 1949.—Egbert H. Walker PLATE 7 1. HOOKER VALLEY BELOW THE HERMITAGE, EAST SIDE OF SOUTHERN ALPS, CANTERBURY PROVINCE, SOUTH ISLAND A debris-choked glacial valley with braided streams thick with glacial grindings is gradually invaded by wild Irishman shrubs (Discaria towmatou) and alluvial fans of weathered rock. (Photograph by E. G. Holt, U. S. Soil Conservation Service.) CS '¢ r - £ aS >” SOUTH ISLAND Rain clouds, rising against the range, drench the rain forests on the lower slopes and cover the higher peaks with deep snow, whence flow the glaciers. (Photograph by E. G. Holt, U. 8. Soil Conservation Service.) Smithsonian Report, 1949.—Egbert H. Walker PLATE 8 1. EROSION AND ITS CAUSE NEAR WAIHO DOWNS, SOUTH OF TIMARU, CANTERBURY PROVINCE, SOUTH ISLAND When sheep overgraze the grass and cut the turf, erosion occurs. Fencing out the sheep (left) allows the vegetation to help control erosion. (Photograph by E. G. Holt, U.S. Soil Conservation Service.) 2. AN OVERGRAZED HILL IN TUSSOCK-GRASS COUNTRY, LINDA PASS, OTAGO PROVINCE, SOUTH ISLAND Soil, bared by overgrazing between the tussocks, is covered with pale green scabweed (Raoulia lutescens) which in turn nurses tussock-grass seedlings. (Photograph by E. H. Walker.) Smithsonian Report, 1949.—Egbert H. Walker PLATE 9 reat Ee 1. PASTURE LAND, HERETAUNGA PLAIN, HAWKES BAY, NORTH ISLAND Rich pastures of introduced grasses are often separated by hedges and windbreaks of lombardy poplars and Monterey pine or cypress. Turnips or swedes are grown for winter feed. (Photograph by E. G. Holt, U.S. Soil Conservation Service.) 2. BREAST HILL STATION IN THE LOW TUSSOCK LAND OF CANTERBURY PROVINCE, SOUTH !JSLAND Sheep stations (ranches) are protected by windbreaks of planted pine. The roadside vegetation is little grazed and more luxuriant. (Photograph by E. G. Holt, U.S. Soil Conservation Service.) Smithsonian Report, 1949.—Egbert H. Walker PEATE LO CANTERBURY PROVINCE, SOUTH ISLAND ” The ‘‘pimples’”’ on the near end are flowers. This weird plant is characteristic of open shingle slopes in the dry area. (Photograph by L. Cockayne, courtesy New York Botanical Garden.) a ac 7 E ee mes - ea i 2. BULL KELP (DURVILLEA ANTARCTICA) ON ROCKS AT LOW TIDE, DOG ISLAND, FOVEAUX STRAIT This flat-bladed leathery kelp grows abundantly on wave-lashed recky shores. (Photograph by L. Cockayne, courtesy New York Botanical Garden.) NEW ZEALAND—EGBERT H. WALKER S20 The kauri forest is the best known of the rain-forest associations, this being due to the noble as well as the highly commercial attributes of its prominent component, the kauri tree itself. (See pl. 2, fig. 1.) Originally almost all of North Island north of the 38th parallel, which crosses the island about the base of the Bay of Plenty, was a vast kauri forest. The title of a recent booklet, “The Waipoua Forest: the Last Virgin Kauri Forest of New Zealand” (21), shows its present state. The Waipoua State Forest or the Waipoua Forest Reserve in North Auckland contains about 40,000 acres, of which about 27,600 acres is actually forested. Perhaps the Trounson Kauri Park with only about 975 acres is too small to consider, but nevertheless it con- tains a primeval kauri stand of limited extent, donated by its lumber- man namesake to the State for a preserve. Because of its small size and the natural degeneration from its exposed margin, its longevity as a primeval forest may be limited. The kauri forest formation is composed of many other tree species besides Agathis australis, as well as characteristic shrubs and ferns, including huge tree ferns, and a few herbaceous plants. Among the many climbers is the kie-kie or climb- ing pandanus (Freycinetia banksit) (pl. 2, fig. 2), the only New Zealand member of this tropical family, Pandanaceae. This forest varies according to the presence or absence of the actually dominant trees, especially of Beilschmeidia taraire in the northern and B. tawa in the southern part of its range. Various studies have been made of this forest from different points of view, but probably Cockayne’s (5) and McGregor’s (21) are of greatest interest to the botanist. Impermanence seems to be woven into the fabric as well as the history of the kauri forest. Cockayne (6) has stated that the kauri is always in a state of progression or retrogression. Seedling kauris cannot normally reach maturity within the kauri forest itself, but must grow up where there is more light, for this forest is predomi- nantly dark, gloomy, dense, and almost impenetrable. Seedlings grow well, however, in the shade of the manuka or in accidental clearings and along roadsides cut through the forest. By some it is thought that the kauri forest will in time cease to exist unless man takes a hand to perpetuate it. But the Maori people first came only 800 years ago and the white man less than 150, and there were kauri forests many centuries earlier. There is in New Zealand now a lively controversy over the preservation of the Waipoua State Forest. Shall it remain as it now stands, or be reduced in size? Shall it be left wholly untouched, or is it to be altered within by forest manage- ment? Shall it furnish timber to commercial exploiters or forest managers, or be a recreation ground and memorial of the past to the 6 Actually the kauri tree is not dominant, as it grows singly or in clusters. Kauri forests are so named because of the prominence and commercial value of the kauri, The broad-leaved dicotylous species are actually dominant. 866591—50——22 328 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 public, or an untouched natural research laboratory to the scientist? Able protagonists have arisen to fight for its preservation from com- mercial exploitation (21). The many complexities and variations in the mixed dicotylous- taxad forest can only be suggested here. Such a forest may be a rimu forest with the ‘‘red pine” (Dacrydium cupressimum) dominant, a totara forest with Podocarpus totara dominant, or a tawa forest with the principal trees Beilschmeidia taraire or B. tawa of the dicotyledon- ous family Lauraceae, though usually with Podocarpus or Dacrydium also present. These forest types intergrade and vary extensively, so it is often difficult to determine just what kind of tropical forest one finds himself in. The parallels of 38° and 42° south latitude are significant as the southern limits of quite a number of species and the northern limits of others. As noted above, the kauri ceases at about 38° south. Some of the 100 or so others which drop out at this point are the toru (Persoonia toru—Proteaceae), the white-flowered tawari (Izerba breri- oides—Saxifragaceae), the climbing fern (Lygodium articulatum), and the trailing fuchsia (Fuchsia procumbens—Onagraceae). The 42d parallel, where many other species drop out, cuts off the northern end of South Island. The fact that this unnatural boundary does not co- incide with Cook Strait between North and South Islands suggests the geologically recent separation of the islands. In addition, certain plant forms drop out at various points. For example, the numerous epiphytes of the mixed rain forests of the west coast of the South Island are conspicuously lacking in the rain forests of Banks Peninsula, the vicinity of Dunedin in Otago, Southland, and Stewart Island. The composition of these mixed rain forests changes also with variations in the atmospheric humidity, the soil moisture and composition, the altitude, and perhaps the geological and biological history. The dicotylous-taxad association of the better-drained land gives way to a much more pure association of ‘‘ white pine,” kahikatea (Podocarpus dacrydioides) in poorly drained or swampy lands, as formerly existed in some prominence in Canterbury on the dry side of South Island. Here the deleterious effect of the drier air, inimical to the best growth of most rain-forest trees, was compensated for by the greater and more steadily available ground water. The swamp- land forests of Stewart Island are dominated by another tree, Dacry- dium intermedia. Altitude, or the changed environmental factors that go along with changes in altitude, also cause changes in this mixed association. As it rises higher on the mountains it is gradually replaced by southern-beech forests. On Mount Egmont, however, they give way to another altitudinal forest association composed of kamahi (Weinmannia racemosa—Cunoniaceae) and the New Zealand NEW ZEALAND—EGBERT H. WALKER 329 cedar (Libocedrus bidwillii—Pinaceae). ‘These numerous changes are dealt with by Cockayne (7) and in many special accounts and reports of the vegetation of certain localities. The southern-beech forest extends intermittently, essentially from the central plateau of North Island to Southland in South Island (pl.1, fig. 1), but is lacking in a stretch of Westland and does not occur on Stewart Island. In former times it was often called birch, but this erroneous designation is seldom used today. These forests are usually composed of just one species of Nothofagus, or at most two. The spe- cies change rather strikingly with changes in altitude. The forester is not content with the taxonomist’s five species of this genus, but is able to recognize in this complex a great many more entities or varia- tions, most of them more or less significant to him in dealing with this economically important forest association. It is now the most im- portant native timber tree, since the kauri is practically gone as a source of commercial timber. Hence, the foresters are working out ways of conserving, extending, and using the beech forests of South Island. Soil conservationists are interested in Nothofagus forests, for these can prevent erosion on mountain slopes, if man would only let them alone or aid Nature to restore them. But man’s ax and fire have laid waste large tracts of mountain slopes. And now another danger threatens the beech forests. ‘The deer which man introduced for sport, or their progeny, are busy browsing and trampling the young beech seedling so that in places the undergrowth of young seedlings has disappeared. The solution for the regeneration of these forests seems to be the extermination of the deer, or at least the keeping of the population under control at a much lower level than it is today. Indeed, the problems of future forests involve many factors in this fascinating country. THE FLORA, OR THE PLANT SPECIES COMPRISING THE VEGETATION Many species of New Zealand plants are possibly unimportant in considering the structure, origin, and distribution of the vegetation or plant associations. They are, however, extremely important in rela- tion to the flora as a whole and the understanding of its origins, affinities, and distribution. There are too many interesting species to deal with them at all thoroughly here, but some aspects of the flora can be reviewed. ‘The marine and fresh-water algae (pl. 10, fig. 2), and the other lower cryptogams must be omitted, but with full recog- nition of their great interest and importance. Although New Zealand is famous as a pteridologist’s paradise, the ferns, too, must be largely ignored. (See pl, 5, fig. 1.) One’s first impulse in looking at the New Zealand flora is to divide it into two categories, the native plants and those introduced by man. 330 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 Each seems a separate unit and the visitor with limited time tends to pass over the introductions. There are, however, so many interesting and important botanical features in both floras that neither should be omitted. THE NATIVE FLORA The indigenous flora of New Zealand is among the most distinctive of the world’s floras. It is not especially rich in number of species, Cheeseman’s Manual (4) listing only 1,763 species, but it contains a wealth of fascinating members, mostly belonging to genera and families unfamiliar to the traveler from northern lands. About 78 percent of the indigenous species of ferns and seed plants are endemics, that is, are not found anywhere else in the world. This becomes 88 percent if the ferns and monocotyledonous plants are excluded, and even higher if only the forest species are considered. Forty genera are found only in New Zealand. Among the more conspicuous or signifi- cant are the lacewood (Hoheria—Malvaceae), one of the few natives which has conspicuous and ornamental flowers and is deciduous, and two composite genera, Haastia and Raoulia, which form the ‘‘ vegetable sheep” (pl. 3, fig. 2, and pl. 10, fig. 1), for which New Zealand is noted. The reasons for the occurrence of these species and genera only in New Zealand are mostly obscure, and leave much yet to learn. They may have evolved here or they may have had a widespread distribu- tion and have died out elsewhere. It cannot readily be determined that these represent a flora which developed in New Zealand, rather than one that came from some other part of the world in remote geological times, but the evidence is that such flora did evolve here. It is called the palaeozelandic flora and certain genera are tentatively assumed to belong to it. They include the three strictly endemic genera just mentioned, besides others now found elsewhere but which probably originated here. Among them are the distinctive New Zealand pine (Dacrydium) in the Taxaceae, the New Zealand broom (Carmichaelia—Leguminosae), the widespread shrub Coprosma (Rubi- aceae) of distinctive divaricating habit and many species, and the woody genus Hebe (Scrophulariaceae), which is often combined with the widespread genus Veronica with mostly herbaceous species. The majority of the nonendemic species and genera are found also in Australia. One might suppose this indicated a relation between the floras of the two regions, but too much emphasis should not be given to this numerical superiority of the Australian element. It is just as important in considering this relationship that certain prominent Australian groups do not occur in New Zealand. Thus in these islands there are no native eucalypts (Hucalyptus—Myrtaceae), bottle- brushes (Callistemon—Myrtaceae), Melaleuca (Myrtaceae), wattles (Acacia—Leguminosae), and other significant genera, especially NEW ZEALAND—EGBERT H. WALKER 331 legumes, and only two genera, Persoonia and Knightia, of the con- spicuously Australian family Proteaceae. Few of the Australian ele- ments are familiar to travelers from the North Temperate Zone. One that a visitor to the New Zealand forest will soon meet, however, is the supplejack (Rhipogonium scandens—Liliaceae), a conspicuous vine which hangs from the tops of the trees. Another is the genus Celmisia (Compositae), whose many New Zealand species are among the chief ornamentals in the montane and alpine vegetation (pl. 3, fig. 2). Besides the Australian element there are indigenous species and genera, found also in the Malay Archipelago and the Pacific region, but they are fewer than the Australian ones. The kauri is of this group, for certain other species of Agathis grow in Australia, New Caledonia, Fiji, and elsewhere. The climbing pandanus (Freycinetia) (pl. 2, fig. 2) and the only New Zealand palm (Rhopalostylis) (pl. 8, fig. 1) belong to definitely Malayan-Pacific groups. The third group of indigenous but not endemic elements is the sub- antarctic, most conspicuously represented by the southern-beech (Nothofagus), already mentioned. There are a good many genera and some species in this group with distribution around the southern Pacific, some with extensions into more northern regions. They are of much interest to plant geographers in understanding the past geo- logical history of this whole southern region. Finally, there are New Zealand plants with a world-wide distribution constituting the cosmopolitan element. An example is the widespread bracken, Pteridium aquilinum, the New Zealand variety of which, var. esculentum, grows to such great size, as already noted. Most of these cosmopolitan species are seashore or littoral plants. These floristic elements and their origin have been discussed by various authors, including Wallace (27) and Cockayne (7). The sub- antarctic flora has been most thoroughly dealt with by Hooker in his introduction to the Flora Antarctica (14) and by Skottsberg (24). BOTANICAL DISTRICTS More significant to the traveler than the origin of the indigenous elements of the flora, is the distribution of the species within the islands. This has already been mentioned in connection with the various plant formations and their occurrence, but consideration of the species rather than the formations brings out more clearly the botanical divisions of New Zealand. Through a lifetime of study of the vegeta- tion and plants of this area, Cockayne (7) has divided the country, exclusive of the outlying islands, into 16 more or less distinct botanical districts. There is, of course, some disagreement with these divisions and other workers will alter them when new studies and interpre- tations are made, as Cockayne himself predicted. The districts are 332 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 shown on the accompanying map, figure 2.” The following char- acterizations are largely derived from Cockayne’s publications and the writer’s personal observations. Only a few of the many outstand- ing species are here mentioned. Northwest of North Cape lies the seldom-visited Three Kings Dis- trict composed of a small group of islands with this name. Their flora includes 10 species of plants not found elsewhere in New Zea- land, 6 of which are obviously related to New Zealand species, 3 have relatives only toward the north, and 1, a species of Chloris (Gramin- eae), is of almost cosmopolitan affinity, although not found elsewhere in New Zealand. The impoverishment of this flora by introduced grazing animals, especially goats, and its recovery after these were removed is an interesting study with significant implications for other likewise devastated areas. These Islands have recently been treated botanically by Oliver (23) and Baylis (8). In the North and South Auckland Districts there are more than 100 species of plants which do not occur farther south, or extend only a short distance beyond the southern limit at approximately the 38th parallel. The kauri tree has already been mentioned in this connec- tion. The taraire (Beilschmeidia taraire), usually the dominant tree in the kauri forest, does not go farther south as does the other impor- tant kauri associate, the tawa (B. tawa). Several ferns are found only here, as well as the parasite Cassytha paniculata (Lauraceae), which resembles, but is no relative of, the dodder (Cuscuta) of wide distribution. This parasite genus occurs in the Pacific area, Australia, and the gumlands of northern North Auckland. It is a hazard to walking wherever it grows as it binds together the manuka and other shrubs by dull green resistant cords. An interesting phenomenon is the occurrence on the small islands adjacent to these districts of species or varieties similar to those on the main island, but with larger leaves, flowers, or fruits, among other differing characters. These islands are difficult to reach, but one can often find these distinct plants growing in private gardens, nurseries, and parks where enthusi- astic New Zealand botanists grow them. Several species have their northern limit in the South Auckland district, one being the silver beech (Nothofagus menziesii). Although there are no natural grass- lands in this area, there are extensive man-made pastures with certain distinct plant species, mostly grasses. The mild climate makes it possible to grow certain citrus fruits and other subtropical crops with success. ¥ After this map was prepared Cockayne and Allan (8) proposed an additional district, the Sounds-Nelson Botanical District, comprising the South Island portion of the Ruahine-Cook District, which formerly straddled the Cook Strait. This district was recognized in Cockayne’s larger work (7), which is followed in this survey, but which does not have a map suitable for reproduction here. NEW ZEALAND—EGBERT H. WALKER 333 THREE KiWGS Is 5° 1 North Cape C. Maria v. Dieman \ Doubtless Bay Mangonui® R Hokianga Hr. at ae 2 Qonegr BARRIER I. | EXPLANATION. C. Colville 2 BOTANICAL (DISTRICT. AUCKLAND : Bay of Plenty r foe eWHITE 1 Three Kings .... North Auckland South Auckland d } East Cape (2) Waikato Subdistrict ay 4, :-(b) Thames Subdistrict NORTH : UA Volcanic Plateau ag a \ Fo East Cape... oobr A: K Berea epesaui es ae \ Me paremoany | Rushine-Cook .... oi @Gisborne ISLAND Y (a) Wellington Subdistrict (b) Marlborough Sounds Subdistrict North-eastern South Island .... North-western South Island ... Eastern South Island Western South Island North Otago .... eee a! South Otago .... =< an Fiord nee = 2 Stewart : Se B. ri Sere} 2457 HANMER oN pba wt ae Greymouth Taramakau R. “runui R. 2 = AWaimakariri R. a @LCHRISTCHURCH Banks Penin. Jackson Hd. ez Tt, xe ingBiigy OR 6 ES en ‘ta R, Big Bay. Sout § u . j, S — Milford Sd.¢ ‘/%*. = et aitaki R. Map of aru ™ NEW ZEALAND L. Te(A nau we rg Aira i showing Sy sy 2a? // DUNEDIN * Tetra ets A 4 Ry AT 42,, Proposed Botanical Districts oy i C teri. R. ; J rs - Tach 00 L t iF, A Invercargill fy, {Ma R. 100 75 50 25 O 1 feces canter or am “BBet Pt, = Pane =, English Miles a ° of Ta R. * STEWART istAND Ey Ss ge 5 L. Cockayna Des. = =n Figure 2.—The botanical districts of New Zealand, according to L. Cockayne (6). 304 ANNUAL REPORT SMITHSONIAN INSTITUTION, 1949 The first natural or tussock grasslands as one progresses southward are found in the Volcanic Plateau District. Here one begins to see certain characteristic subalpine shrub formations, especially of monoao (Dracophyllum subulatum—Epacridaceae). The volcanic mountains, three of which are still active, raise their summits far above timber line and support a fascinating alpine flora. The northern tip of the East Cape Botanical District is within the northern plant zone marked off by the 38th parallel, so contains some northern plants. The southern part in the Hawkes Bay region is drier, hence it has certain agricultural possibilities not found else- where. Maize or Indian corn (Zea mays—Gramineae) is grown for its grain, but elsewhere mostly for fodder. To the botanist Mount Egmont is probably the first attraction of the Egmont-Wanganui District in the southwest portion of North Island. However, its high mountain flora is less rich than is that in the mountains of the central plateau, which are visible in good weather from Egmont’s higher slopes. The zones of vegetation on this isolated volcanic cone, however, are very vivid. There are few plants endemic to this district and the southern-beech forests are absent.