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George Henry Falkiner Nuttall and the Nuttall Tick Catalogue

by James E. Keirans1/

From the turn of the century until his death in 1937, George H. F. Nuttall amassed what at the time was the world's largest tick collection. This collection was the basis for monographic revisions of the genera Argas and Ornithodoros (1908), Ixodes (1911), Haemaphysalis (1915), and Amblyomma (1926). These works are still consulted by specialists and, although outdated, the Amblyomma revision remains the best published work on that genus.

Nuttall was in a unique position with regard to the collection of ticks. He personally collected, but that was a secondary source of material. As Director of the Molteno Institute for Research in Parasitology, Cambridge University, he had access to collections by present and former students and corresponded with parasitologists throughout the world. He also exchanged tick specimens with most of the leading tick workers of the day, including L. G. Neumann, W. Dönitz, H. de B. Aragão, E. Brumpt, and R. A. Cooley. Readers of the scientific publications "Parasitology" and the "Journal of Hygiene," both of which he edited, were acquainted with his numerous publications on ticks and sent him samples for identification. More importantly, this was the era of the greatest expansion of the British Colonial Empire and the time of the formation of the Entomological Research Committee (Tropical Africa), later expanded to the Imperial

Bureau of Entomology and now the Commonwealth Institute of Entomology. This committee began the first systematic collection and study of insects and other arthropods injurious to man, animals, and crops in Africa (Anon., 1909; Shipley, 1910). Nuttall was a member of this committee and was responsible for all tick identifications.

However, all this access to huge tick collections would have been in vain had it not been for Nuttall's meticulous recordkeeping. Each collection received a Nuttall number, and all data were put on labels kept with the ticks in alcohol; numerically, the data were entered in his tick catalogue. The large number of ticks from collectors throughout the world, combined with Nuttall's detailed accurate recordkeeping and his taxonomic expertise, led to a tick collection unrivaled in its day.

After Nuttall's death in 1937, Cambriage University presented his tick collection to the British Museum (Nat. Hist.) with the single stipulation that it remain a separate entity as--The Nuttall Tick Collection--and not be incorporated into the general collection. Various investigators have studied parts of Nuttall's collection when they pertained to species groups in which they had an interest, but the entire collection has never been revised and updated. Since his death, taxonomic concepts have changed within the Ixodoidea and their numerous vertebrate hosts. In addition, the emergence of scores of new nations, especially in the post-World War II period, requires an updating of all locality data associated with the Nuttall collection.

1/ Research entomologist (medical), Department of Health and Human Services (HHS), Public Health Service, National Institutes of Health, National Institute of Allergy and Infectious Diseases, Epidemiology Branch, Rocky Mountain Laboratories, Hamilton, Mont. 59840; now in HHS, Entomology Department, National Museum of Natural History, Museum Support Center, Smithsonian Institution, Washington, D.C. 20560.

Biography of George H. F. Nuttall

George Henry Falkiner Nuttall was born in San Francisco, Calif., on July 5, 1862, the second of three sons and two daughters of Robert Kennedy Nuttall, M.D., and his wife, Magdalena. His father (1815-81) received his doctor of medicine degree from Aberdeen in 1847 and was a physician in the prison colony in Tasmania for 4 years before migrating to San Francisco and establishing an obstetrical practice. In 1865, the family moved to Europe, where the children were educated in France, Germany, England, and Switzerland. In 1873, his father took a house in Richmond, Surrey, and George went to Epsom for a part of his primary school education; in 1876, he was privately tutored before returning to America in 1878. It was to this cosmopolitan upbringing that Nuttall owed his ability to speak German, French, Italian, and Spanish, accomplishments that were to help him greatly in his scientific career and travels. He entered the University of California, Berkeley, to study medicine and obtained a doctor of medicine degree in 1884 and also won the Kane Prize.

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After 1 year at Johns Hopkins University in Baltimore, Md., Nuttall went to Germany--first, to Breslau and later, Göttingen, where he became interested in parasitology, which became the main scientific interest of his life. 1891, he returned to America as assistant to W. H. Welch, professor of pathology at Johns Hopkins. From 1892 to 1899, he worked on hygiene at Göttingen and Berlin. In 1895, he married Paula von Oertzen-Kittendorf (1873-1922) of Mecklenburg. They had two sons, George (b. 1896) and Winfred (1897-1972), and a daughter, Carmelita (b. 1902). In 1899, Nuttall gave lectures on bacteriology at Cambridge. A year later, he was appointed university lecturer in bacteriology and preventive medicine.

In 1901, Nuttall founded the "Journal of Hygiene," which he edited up to the time. of his death. Seven years later he began "Parasitology," which he edited until 1933. The following note appears in

Nuttall's journal for May 1900: "In this month I saw Dr. John Haldane, F.R.S., at a meeting of the Physiological Society in Cambridge. We had a conversation which led to our starting the JOURNAL OF HYGIENE the public announcement of which appeared in October, the first number coming out in January 1901. The suggestion initiated from Haldane, who being a physiologist was unwilling to take on the chief editorship. Dr. Arthur Newsholme, representing the administration side of Hygiene, joined us as third editor and we soon secured a lengthy list of collaborators in all parts of the world." In his editorial work he displayed the same. thoroughness as in his research. He considered that part of the duty of an editor was educational and spent much time in correcting and improving papers and in advising young and inexperienced workers. As editor, he exerted great influence on investigators all over the world, and his journals became models upon which the publications of several scientific societies were based.

Among his outstanding contributions to science was his discovery while working in Flugge's Institute (1888) that defibrinated blood possesses a strong bactericidal property against anthrax bacilli and that this property disappears by heating the blood to 55° C. The results of this work were incorporated in a paper "A Contribution to the Study of Immunity," which received the triennial Boylston Prize from Harvard University. This work initiated the study of humoral immunity and was the forerunner of such great discoveries as antitoxic immunity by workers like E. A. Behring and S. Kitasato. It occupied also a prominent place in the discussion of humoral versus cellular immunity formulated by E. Metchnikoff.

In collaboration with W. H. Welch (1892), he studied in great detail the anaerobic gas-producing micro-organism known now as Clostridium perfringens, the agent of gas gangrene, the importance of which as a pathogenic agent was not fully appreciated until World War I.

In collaboration with H. Thierfelder (1895-97), he carried out the first successful experiments on life under aseptic conditions. In these investigations, guinea pigs delivered by Caesarean section were kept alive and fed in a specially devised chamber under bacteria-free conditions.

In 1897, Nuttall turned his attention to the role of arthropods in the spread of disease, a subject to which he devoted his later life. In 1899, he published his well-known paper "On the Role of Insects, Arachnids and Myriapods as Carriers in the Spread of Bacterial and Parasitic Diseases of Man and Animals," which contains an exhaustive critical and historical review of the entire subject. Soon afterward (1900-01), he became interested in the history of malaria in England and in collaboration with L. Cobbett, T. Strangeways, and others undertook a survey of the distribution of the genus Anopheles in England. He found that three species of these mosquitoes were in all districts that were formerly malarious. The disappearance of malaria in England was therefore not due to the extinction of Anopheles. In collaboration with A. E. Shipley, he carried out an important investigation on the structure and biology of Anopheles, which is still considered a classic.

About the same time (1901), Nuttall turned his interest to precipitin reactions. He devoted the next 3 years to studying the application of the precipitin reaction to phylogenetic relationships among vertebrates. In 1904, he published his classic monograph "Blood Immunity and Blood Relationship," incorporating the results of an extensive investigation of precipitin tests carried out on the blood of about 600 species of animals. This investigation clearly established the correlation between the antigenic relationships of animal sera and the zoological relationships of the species. It threw a new light on the study of the phylogenetic relationships in the animal kingdom and provided also a new method for identifying the character of minute

traces of blood, a procedure that has had notable applications in forensic medicine.

After the publication of this monograph, Nuttall began his investigations on diseases transmitted to animals by ticks. By means of infected ticks (Haemaphysalis leachi) sent to him by C. P. Lounsbury from South Africa, he succeeded in infecting dogs with the agent of piroplasmosis, a disease unknown in England. This was the first case of investigating a disease imported by an infected vector. An important result of this study, which had great economic importance, was the discovery of the curative property of trypan blue for piroplasmosis in dogs, cattle, and sheep. This study was followed by an extensive investigation of the anatomy, biology, life history, and systematics of ticks, carried out in collaboration with C. Warburton and L. E. Robinson. The results of these investigations are incorporated in numerous papers and in the monographic study of ticks that occupied him on and off during the remaining years of his life. During this investigation Nuttall acquired a very large collection of ticks from all parts of the world, at that time the largest in existence and rich in type specimens.

Most of his research interest during and after World War I was in the study of the Anoplura, their life history, biology, anatomy, and the problems of combating the spread of these parasites.

In his early life Nuttall had traveled in Mexico, Cuba, and North America, and it was probably during this period that he became interested in natural science. His uncle, Tiburcio Parrott, gave him his first microscope and started him in making and preparing microscopic slide mounts of all varieties of animals and plants. This was perhaps the initial event in Nuttall's scientific career.

Nuttall's scientific work shows that he possessed the attributes of the acute observer, the experimental scientist, and the naturalist to a remarkable

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As a naturalist, he showed a real passion for collecting all sorts of specimens and in preserving, classifying, and cataloguing them. This work was always carried out with a great regard for permanence. He selected always the correct container for the specimen, unfading ink, and paper of lasting quality. The catalogues of his various collections have several hundreds or, in the case of ticks, thousands of entries in his own handwriting, giving the complete history of specimens to which they refer. In his journal, with equal meticulousness, Nuttall recounted his daily experiences, conversations, trips to various meetings, and interesting visitors to his laboratory and home.

In addition to his scientific attainments, Nuttall read widely on a variety of subjects and spoke with expertise on such topics as heraldry and gardening. He was also an excellent draftsman with a great interest in art. In 1904, he was elected a Fellow of the Royal Society, at the same time as his friend A. E. Shipley, and the year 1906 was one of the most important of his life as shown by the following journal entry.

"July 20 admitted to a Junior Fellowship at Christ's College. October 16 elected Quick Professor of Biology at Cambridge, and duly admitted on the 19th by the Vice-Chancellor (Mr. Roberts of Caius) at the Lodge and on the same day went to Brighton to see Mother back to C. 24th. 24 May proceeded to the degree of Dr. of Science (Cambridge). November 20 dined with Prof. Newton, F.R.S., at Magdalene College after having at 11 a.m. been one of a deputation which was presented to the King of Norway at Buckingham Palace on behalf of the Royal Institute of Public Health.

"This was a most eventful year. Following upon the San Francisco disaster which for some time we thought might end in the loss of our fortune, things began to look up and in the end the fireinsurance companies paid all that was due, less 5% for cash payment rec'd within a year of the disaster.

"It was a great satisfaction to be admitted a Fellow at Christ's for I felt that I had at last got into 'the inner circle.' Soon after, I was made Reader in Hygiene, the Lectureship in Bacteriology and Preventive Medicine, previously held being allowed to lapse. Curiously enough I only held this post for 4 days before I was elected to the newly established Quick chair when, in turn, the Readership lapsed, for both the Lectureship and Readership had been personal appointments. When the Readership was established, there had been no thought of my going in for the Professorship, for the conditions under which the latter was to be held had not as yet been determined. However, as time went on, it was decided that the holder of the chair should devote himself to the 'study of the protozoa, especially in relation to disease,' and this being the case I made an application for the post. The decision as to who should be the first holder of the chair was arrived at with some difficulty and a good deal of heart burning. My friends rallied nobly to the charge and I was elected. It is just as well to forget the disagreeable aspects of the campaign for it ended in Victory for me. In my printed application with the friend much matter relating to this event, so important for me & mine for it put me 'upon my legs' again after the San Francisco disaster which strained our finances badly, my income from that source having been cut off for upwards of a year.

Nuttall received honorary degrees from the Universities of California, Strasburg, Egypt, South Africa, and Liège and was elected corresponding member of many scientific societies. He received also the Belgian Order of Leopold II and the

French Order of Commandeur de la Légion d'Honneur.

On the afternoon of December 16, 1937, Nuttall rushed from Cambridge to London to attend an autograph party at Foyles Bookstore for his son, George, who had coauthored a book with H. M. Hyde on air defense and the civil population. From there he dashed off to an art gallery where the portraits of Philip Alexius Laszlo de Lombos were on display, including one of Nuttall painted in 1932 (see frontispiece). From there he went immediately to the Craven Hotel in Charing Cross where he suffered a fatal heart attack. Unfortunately, it was on the eve of a dinner in his honor to be given by 60 colleagues upon his retirement from his long and successful editorship of the "Journal of Hygiene."

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