Protist, Vol. 152, 6985, May 2001 Urban & Fischer Verlag http://www.urbanfischer.

de/journals/protist

Protist

FROM THE ARCHIVES

Two Most Remarkable Amoeba Men: Joseph Leidy (18231891) of Philadelphia and Eugne Penard (18551954) of Geneva
John O. Corliss1
P.O. Box 2729, Bala Cynwyd, PA 19004, USA

How can life be tiresome so long as there is still a new rhizopod undescribed? Joseph Leidy Joseph Leidy and Eugne Penard, early pioneers and experts without peer (in their times and still today) in detailed study of the morphology and taxonomy of the free-living rhizopod Sarcodina, shared possession of a great curiosity about and a loving devotion to these fascinating if oft-neglected protists. But, otherwise, both the professional and personal lives of these two individuals differed quite markedly. As we shall see, our pair of distinguished gentlemen had little in common beyond their passion and ability to carry out invaluable researches at the microscopical amoeba level, no easy accomplishment in the times of their work. Our debt to them as outstanding biological scientists will endure forever, although as dedicated naturalists they never achieved the fame and recognition accorded the inspired experimentalists and conceptual theorists of their day and of all times since then and, alas, they never shall. A further factor, which they share in being largely neglected today, is their lack of establishment of a school to follow in their scientific footsteps: both men were essentially loners, having neither students nor close colleagues as joint-authors. Thus the present article is offered, in part, as a tribute to all such dedicated fact-finding workers of the past widely forgotten, today, following advent of more sophisticated (and often experimental) ap1

proaches to solution of many of our age-old biological problems.

Brief Historical Background

In the broad area of the systematics of species belonging to major groupings of the Sarcodina sensu lato (but mostly excluding, here purposely reference to fossil and parasitic forms and most members of the Radiolaria), other principal workers predecessors or contemporaries of Leidy and Penard of the period stretching from early nineteenth to early twentieth century deserve at least brief mention, with specific citation to several of their main publications: Archer (numerous scattered papers, not cited here, period 18661880), Awerinzew (1906), de Bary (1859 et seq.), Blochmann (1886), Btschli (18801882), Carter (scattered works, not cited, period 18561870), Cash and Hopkinson (1905, 1909), Cash and Wailes (1919, 1921), Cash et al. (1915), Cienkowski (1863 et seq.), Claparde and Lachmann (1858-1861), Dobell (1919), Dujardin (1841), Ehrenberg (1838), Greef (scattered papers, not cited, period 18661879), Haeckel (1878, 1894), Hertwig (1879), Hertwig and Lesser (1874), A. and G. Lister (1925), J. Mller (1858), Perty (1852, and earlier), Rhumbler (1903), Schaeffer (1916, 1920, 1926), Schewiakoff (1926), Schulze (1874), Wailes (1912, 1913), and Wallich (scattered papers, not cited, years 18631875).
1434-4610/01/152/01-069 $ 15.00/0

fax 1-610-664-4904 e-mail jocchezmoi@aol.com

70

J. O. Corliss

From the above-mentioned 100-year period until nearly the present, additional protistologists contributed heavily to further advances in understanding better the taxonomy of diverse sarcodinid taxa: Alexopoulos, Bardele, Bonnet, Bovee, the Cachons, Chardez, Chatton, Cushman, Declotre, Deflandre, De Saedeleer, Febvre, Febvre-Chevalier, Foissner, Grell, Grospietsch, Haynes, Heal, Hedley, Hollande, Hoogenraad, Jahn, Jepps, Jung, Kudo, Khn, Lee, Loeblich, Ogden, Olive, Page, Patterson, Raper, Sawyer, Schnborn, Singh, Tappan, Trgouboff, Valkanov, and Vickerman: see references in Bovee and Jahn (1973), Corliss (19781979, 1992), and contemporary protistological compendia such as Lee et al. (1985, 2001) and Margulis et al. (1990). Seldom, however, are the publications of the above researchers of the monographic proportions of some of those cited in the preceding paragraph, although the prodigious output of excellent systematic and descriptive papers and reviews by three modern workers E. C. Bovee, K. G. Grell, and F. C. Page warrant special recognition with regard to the rhizopod amoebae overall. A bit earlier, a prolific expert on the thecamoebae was G. Deflandre (see citations in Deflandre 1953). Today, as we enter the 21st century, there are surely even fewer protistologists intensively devoting major research effort to amoeba morphology and taxonomy than the scant one dozen of 2530 years ago, a figure that Bovee (in Bovee and Jahn 1973) was justifiably lamenting as too low a number. Incidentally, it is rather surprising that the amazingly perceptive first observer of little things, A. Leeuwenhoek of Delft, never detected or described an amoeba (except for one foraminiferan found in the stomach of a shrimp: see Corliss 1975) amongst the extensive collections of protists he examined with his glorified hand lens in the late 1600s. It was well into the next century that Joblot (1718) gave us a very brief description of what we now know as the heliozoon Actinophrys sol and that Rsel von Rosenhof (1755) published a paper on his celebrated little Proteus; and then still some 30 years after that before the astute observations of O. F. Mller (1786) became available, a monograph including descriptions of a (but very) few amoebae along with those of many other microscopic forms of life.

Figure 1. Joseph Leidy, M.D. (18231891), in his midsixties (a couple of years before his demise). [Photograph from the authors private collection]

Early Life and Education of Leidy and Penard

Joseph Leidy (Fig. 1) was born in the heart of Philadelphia (PA, USA) on 9 September 1823. Like his father and grandfather before him, he was a

true-blue Philadelphian and seldom traveled far from his beloved city throughout his lifetime. His great grandfather, John Jacob Leydig (as the surname was originally spelled in Germany), had emigrated from Wittenberg to escape religious persecution, arriving in Philadelphia in 1729 and settling on a tract of 400 acres purchased from the well known Quaker William Penn. John Jacob, interestingly enough, was a brother of the great grandfather of Franz Leydig [18211905], distinguished histologist and comparative anatomist of Wrzburg and Bonn and a contemporary of Joseph, our American naturalist. And John Jacobs wife was a sister of Francis Joseph LeFebre, a marshal of Napoleon I and a peer of France. Joseph Leidys mother died two years after giving him birth; his father (a hatter by trade) soon (re)married, a cousin of his wifes; and it is to his highly intelligent, cultured, and principled stepmother (Christiana Mellick) that Leidy himself credits his stimulating early home environment. Although German, a common language in various parts of Pennsylvania in those days, was spoken in the home, Christiana insisted on everyones practicing English as well; and, at her instigation, young Joseph, at age eight,

Two Most Remarkable Amoeba Men

71

was sent to a private day school where he began to learn while also taking other studies Latin, Greek, and French, as well as English, all languages to stand him in good stead in later years. Joseph was an able student but preferred studying nature directly rather than undergoing formal education in a regular classroom setting. His avid curiosity about plants, animals, and minerals, combined with a natural talent in artistry, led to production (by age 10 and beyond) of notebooks filled with accurate sketches of rocks and organisms encountered in the wild, with the latter often labeled with their correct Latin names. On days and hours not required to be in the schoolhouse, he preferred such exciting pursuits over playing sports with the other lads, or even getting home in time for dinner! At this stage of his early life, Josephs parents differed concerning what career he should pursue. His father favored art, with heavy emphasis on the practical profession of sign-painting; his (step)mother considered a medical career more appropriate. Fortunately, his mothers view prevailed. At the tender age of 17, Joseph enrolled in the medical school of the University of Pennsylvania, whose buildings of that time were located not far from his home, and received his M. D. degree in 1844 (with a thesis entitled The Comparative Anatomy of the Eye of Vertebrated Animals). His medical training cost his parents some $140 per year but, as a quite comfortably living middle-class family, they managed to make the payments (yet they might well have been distressed if they could have predicted that, only two years out of his M. D., young Joseph would be completely abandoning any active practice of medicine!). While doing not badly in his studies, especially in preparing anatomical dissections and in producing highly accurate drawings, Joseph Leidy still voraciously pursued his own interests as a mostly self-taught naturalist, enhanced by his having been introduced to an unusually good microscope by one of his perceptive mentors and soon thereafter given one as a birthday present by his equally perceptive mother. This instrument allowed him to note and describe cytological and histological details not stressed in his formal course work and, of course, opened the way for his subsequent protistological revelations, to which we return below. In fact, Leidy was the first American to seriously and extensively apply microscopy in biological and medical research. It was also while he was a medical student that he first met the famous British geologist Charles Lyell [17971875] when the latter gave some lectures in Philadelphia; the two became instant friends and mutual supporters (of each other and of Charles Darwin) and got together whenever both were in either America or England at the same time.

Figure 2. Eugne Penard, D.Sc. (18551954), in his late nineties (ca. three years before his passing). [Photograph from the authors private collection]

Eugne Penard (Fig. 2) was born in Geneva, in the predominantly French-speaking section of Switzerland, on 16 September 1855; and like several generations of his direct ancestry he was a patriotic citoyen gnevois throughout his lifetime. Unlike his fellow amoeba man Joseph Leidy, however, he was more frequently absent from his home town and for longer periods of time. The first Monsieur Penard (that we know about) established himself in Geneva back in the middle of the 14th century. This man and a number of his descendants were watchmakers and engravers, not too unusual a profession for Swiss men possessed of the requisite ability to do such precise work. Eugnes adventurous grandfather left his wife and children (in Switzerland) to pursue such a career in America; but his father was director of a private school in Geneva and did not follow in the footsteps of his own errant papa. We know little about Eugnes early childhood, but he studied at the Collge de Genve and, at age 17, accepted a job as a cashier in a bank. Puzzlingly (to me), he remained in that profession for nearly nine years. But, apparently, he and his parents eventually realized that he had abilities (latent and other-

72

J. O. Corliss

wise) beyond such mundane work. He next went abroad for a year or so, renewing his scholastic pursuits, first at the University of Edinburgh, in faraway Scotland, and then at Heidelberg (in Otto Btschlis laboratory), in nearby Germany. His studies, at which he excelled, included among other subjects as well Latin, Greek, Greek Philosophy, Geology, and French Literature. He had already mastered the French, German, Italian, and English tongues; and he was soon to add Russian to his linguistic repertoire. Returning to his home country in 1882, Penard became a zoological student of Karl Vogt [18171895], erstwhile Professor at Giessen, Germany, a position from which he was none too gently removed for having participated in the revolutionary movement of the late 1840s. Embittered, Vogt was happy to have been offered a professorship by the more enlightened University of Geneva, and Penard was an eager and able student. In 1887, the two spent several months at the rather newly founded Stazione Zoologica di Napoli, where Eugne working at the Swiss table in the laboratory completed an elegant investigation of sperm penetration into the sea urchin egg (a work, regretfully, never published). In the summer of the same year, he studied the dinoflagellate Ceratium hirundinella, submitting his results of that research as a (prize-winning) thesis to the University of Geneva, for which he was awarded his Doctor of Sciences degree (in late 1887). We shall return in a subsequent section to Penards fascinating stays in Russia during the years 1883-1886 and 1892-1898, periods not covered above. Suffice it to mention here that, by his own words, he spent not more than five minutes in making original microscopical observations essentially during his whole times there, due to the pressure of his official duties. Can the reader imagine what sorts of jobs could possibly completely isolate him from the pursuits with which he was destined to personally fall so deeply in love, commencing earnestly in 1887 but soon only five years later to be interrupted again until nearly 1899?

Early Professional Work and First Protistological Publications

Tracing exact parallels in the developing lives of the two great amoeba men being briefly recalled in this historical essay is rather difficult though I believe worthwhile for at least five reasons: Leidy was born a whole generation (32 years!) before Penard; Penards life spanned nearly 100 years, Leidys only

68; Leidy entered the microscopic world much earlier (age < 20, while Penards was close to 30); Leidys first substantial protistological publications (occurring long after and/or amongst papers in other vastly different research fields) appeared when he was still scarcely 30 years old, while Penard was several years older than that when his first (the dinoflagellate) monograph was completed, and he was essentially carrying out research solely in the field of protistology; and, finally, Leidy published papers involving description of protists up to and even beyond the year of his death, while poor eye-troubled Penard under doctors orders was obliged to totally give up use of the microscope at age 67: what a blow to a taxonomic protistologist, although he lived on, in excellent health otherwise, until well into his 99th year. Joseph Leidy was precocious both in becoming known and respected professionally on an international level and in producing diverse works of medical and broad zoological interest. By the time he had reached the age of 25, he had already published well over 30 papers, many as very brief notes in the Proceedings of the Academy of Natural Sciences of Philadelphia, but others were several pages in length and accompanied by excellent figures. The amazing range of topics covered ran from the microanatomy of the human larynx and liver and tissue transplantation experiments to description of new genera and species of fossil mammals (including, among others, two rhinoceroses and the early American horse), from the taxonomy of several new species of invertebrates to careful description of their helminth, fungal, and even some bacterial parasites, and from observations on (and superb illustrations of) the internal organs of snails and the nematocysts of Hydra to the crystalline bodies in tissues of certain plants! And works of a purely protistological nature were not long in forthcoming. In fact, by 1850, the same year in which he laboriously edited an American edition of the human anatomy text by the Britisher James Quain, M.D. (Leidy wrote his own, far superior, text on the subject in 1861, with a greatly enlarged and revised 2nd edition in 1889), he was describing new species of symbiotic protists among the gregarine sporozoa and the nyctotheran ciliates. Leidys habit of working six or even seven days a week, and sometimes 20(!) hours a day, made all such accomplishments possible. Yet this highly focused workaholic with an encyclopedic mind also always found time to help other people, personally answering (by long-hand, of course) many hundreds of letters from folk with queries about all sorts of natural phenomena, unstintingly cooperating in es-

Two Most Remarkable Amoeba Men

73

tablishing new Philadelphian institutions or academic departments that included emphasis on basic biological subjects, and holding interesting conversations on the street with both uneducated fishmongers (and identifying some of their unusual catches for them in their market stalls) and University administrators or high-society industrialists and their wives, all the while (appearing, himself, to be) a relaxed, cheerful, kind-hearted, leisurely person. In 1853 (but submitted in finished form to the publisher in 1851), Leidys succinct 68-page publication, A Flora and Fauna within Living Animals, practically established his reputation as the Father of American Parasitology even though he was not yet 30 years of age. Beautifully illustrated by 10 plates of precise figures, the text went beyond the usual morphological descriptions, including life cycle information and data on host-parasite interrelationships as well. Leidy also supplied convincing proof of the falsity of the doctrine of abiogenesis or spontaneous generation (although generally only the well publicized later observations and postulations of Tyndall, Pasteur, Koch, and other Europeans are credited today with such a significant discovery. The French protozoologist Joblot and the Italian physiologist Spallanzani, however, had also demonstrated it in the 18th century: see Cole 1926). Even earlier than 1853, Leidy had observed the very important nematode worm Trichinella spiralis as a parasite in pigs, and he correctly suspected that it might cause the disease trichinellosis (still today widely but mistakenly called trichinosis, although it was legitimately so known in earlier times) in humans if infected pork was insufficiently cooked before being eaten. Eugne Penard, delayed in starting his protistological career until about 33 years of age, certainly made up for lost time. He produced major papers on the heliozoa in 1889 and 1890, following his doctoral thesis on Ceratium, and very soon had published his first comprehensive (230-page) monograph on the rhizopod amoebae of fresh water (Penard 1890c) and a sizable work on the dinoflagellates of Lake Geneva (Penard 1891a); not much later (1893), his paper on the lobosean amoeba Pelomyxa. In 1891, by coincidence the very year of Leidys passing, Penard published a paper in the American Naturalist, in English, on his findings of testaceous or thecate rhizopods, some new to science, in the Rocky Mountains of Colorado, which he explored personally for several months during a visit to America (see below). As he told me with a certain amount of unconcealed delight when I visited him, some six decades later in Geneva, he had climbed the Rockies to 12,500 feet for his highest collection site, bet-

tering Leidys record by some 2,500 feet! He was a great admirer of Leidy, but his own descriptions were even more detailed and he was less cautious about describing new genera and species. He also paid more attention to the rules of the international zoological nomenclature, although, in all fairness to the American, there were none in force at all until 1842, when they were highly voluntary; and the forerunner of the present obligatory Code (the ICZN) did not come into existence until 1889 (see Ride 1999). Penards figures were painstakingly precise, no doubt reflecting an ability inherited from watchmaking forebears, but lacked some of the artistry revealed in the Philadelphians magnificent colored plates. At age 96, he apologized to me for the often sketchiness of his own line drawings in many of his monographs; but they were always exceedingly accurate and were almost always made from living, unfixed organisms. Furthermore, a naturalist endowed with artistic talent, Penard also produced drawings of scenic landscapes, especially of unusual sights during trips away from home. Im pleased to be possessor of one of these, a view of Gold Hill, Colorado. Penards reputation was well established through his early publications, but was even further enhanced as his protistological career continued apace (after another hiatus of some six years abroad: see below). Like Leidy, he remained a modest man while obviously a person of great energy, drive, skill, and dedication to any research project that he chose to pursue. Never a user of the oil immersion lens on his monocular microscope, his eyesight must have indeed approached that of his renowned predecessor Leeuwenhoek. Attesting to that is his early detection, 115 years ago, of the true nature of the second flagellum in dinoflagellates; and, some 80 years ago, of the presence of the reduced second flagellum of Astasia and of the delicate caudal cilium of certain ciliates, the latter organelle sometimes today practically requiring phase or even electron microscopy to clearly demonstrate its existence.

Travels Away from Home Bases

Joseph Leidy, at a time in Americas scientific growth when such scholastically talented youths as he were typically sent off to the great European universities for proper training in their research specialties, quietly chose to remain in Philadelphia. His aim in life seemed to have been to discover and describe (new) biological facts with precision (and enjoyment!), while well aware of the relevant informa-

74

J. O. Corliss

tion available in the burgeoning European literature, and not to worry about speculation, competition, or fame. As he bluntly (rather stubbornly?) stated in the preface of one of his paleontological monographs (Leidy 1869), The present work is a record of facts No attempt has been made at generalizations or theories which might attract the momentary attention or admiration of the scientific community. Yet so voluminous and accurate were his observations that other great 19th century biologists of the world were soon getting in touch with him for further data, many of which supported their own ideas: an outstanding example was Charles Darwin (others, in due time, included renowned paleontologists, human anatomists, and parasitologists). Nevertheless, Leidy but usually only after strong urgings of colleagues did venture outside Philadelphia on occasion: to other parts of the ever-expanding USA and overseas. At the age of 25, he accompanied his University of Pennsylvania Medical School mentor Dr. William Horner to Europe for a profitable period of several months (spring and summer of 1848). His name and reputation (already!) had preceded him and he was cordially greeted by such eminent (or soon to become eminent) scientists as among still others Charles Darwin, Charles Lyell, Richard Owen, and John Tyndall in England; Claude Bernard, H. MilneEdwards, and C.-E. Blanchard in France; K. M. Diesing, Johannes Mller, C. T. E. von Siebold, R. A. von Klliker, and T. Bischoff in Germany; and Karl Rokitansky in Austria. His and Horners scientific and social calendar on the trip was hardly that of the ordinary tourist! They visited innumerable museums, colleges, herbaria, botanical gardens, hospitals, medical research laboratories, libraries, government buildings (e.g., Houses of Parliament), shops, bookstores, and microscope companies as well as art galleries, castles, cathedrals, churches, countrysides, parks, operas, ballets, playhouses, and theatres, not to mention residing in elite hotels and dining in the best of restaurants. Their days and nights were fully occupied through the beneficent arrangements made by new-found friends and hosts. Two years later (in the summer of 1850), persuaded to accompany Dr. George B. Wood this time, Leidy returned to Europe, officially mostly to collect specimens, wax models, microscopes, charts, drawings, and books for medical school teaching needs. The trip was of slightly shorter duration but still covered some of the same activities (including meeting Owen and others again) and was a strenuous one; but Joseph returned in a better state of health than when he had set out on the voyage. His overseas colleagues were obviously im-

pressed by his top-quality research productivity, in a diversity of fields, during the interim twixt his two visits. One can imagine the thoughts that ran through many of their distinguished gray heads: how amazing for an American, and a mere lad, still, at that! Leidy was too busy with fossils in 1858 to cross the Atlantic to carry out another planned trip abroad, so it wasnt until 1875 that he journeyed again to London and Paris (with touristy side trips to Geneva, Brussels, and major cities in Germany), this time accompanied by his delighted wife Anna. Nonetheless, he again managed to make many professional visits and meet old acquaintances, plus acquiring new friends in high scientific positions. One of the major highlights for him was a long meeting with another admirer of his, T. H. Huxley, the great British naturalist, part-time protozoologist, and evolutionary biologist (well known as Darwins bulldog). Leidy, still interested in gems and rocks, also visited a number of distinguished geologists and mineralogists. At their request, he later arranged to have sent to the British Museum several shipments of invaluable specimens from the American Western Territories. His final brief trip abroad, accompanied by his faithful wife and his loving (adopted) daughter Allwina, was in 1889, a scant two years before his death. He was in rather poor health himself, and then aboard ship poor Anna came down with typhoid fever, which obliged them all to spend a number of weeks in London, with husband and daughter literally hovering anxiously over her sick bed. Fortunately, receiving the best medical care available, Anna recovered and they later journeyed on to Paris. Joseph, now a rather tired old man, again brought back to Philadelphia specimens, etc. from various museums; but the whole excursion ended up being an exhausting one. Although Leidy, the Father of American Paleontology, described from tons of material specimens representing more than 300 important new species of animals ranging from invertebrates to fishes, dinosaurs, and mammals, he seldom collected the fossils in person. Bones, teeth, scales, etc. were sent to him by eager collectors from all over the United States. However, he did make a few trips himself, most notable ones to the American far west in the summers of 1872, 1873, 1877, and along the eastern coast (far up into Canada) in 1874. He also, everywhere, collected Indian artifacts, minerals, plant specimens, insects and other invertebrates, parasitic material, and rhizopods! See the following section for more information on Leidys paleontological contributions.

Two Most Remarkable Amoeba Men

75

Eugne Penard was much more adventurous with respect to roaming away from his comfortable residence, in Geneva. In fact, during most of the years 18811898, he was living in other countries. As mentioned above, he studied for a while (in 18811882) outside Switzerland in England and Germany after his bank job; and then, a year or so later, he embarked on his first Russian adventure. He had applied for the position of tutor (perhaps encouraged by his schoolmaster father?) to the royal family of Prince Orloff; accepted, he spent most of the years 18831886 in that busy position. In 1884, he managed to get in a trip to Algiers (perhaps with his Russian employer?). Penard was then 29 years old; and, by chance, he fell into animated conversation with the citys archivist/librarian while searching for certain scientific books (biological?). Quite likely (I wish we knew for certain!) the matter of protozoa arose, although young Eugnes doctoral researches were still three to four years from realization. At any rate, the 42-year old librarian eventually bade him bon voyage and the pleasant encounter ended. Years later, Penard discovered that the friendly old man had been none other than Emile Maupas, the French protistologist who, at the very time of their meeting, was deep into the magnificent ciliate researches being carried out after work in a little laboratory in his small Algerian apartment that would bring him long-lasting fame (e.g., Maupas 1888, 1889; and see Jennings 1929, and the brief accounts in Corliss 19781979 and Nanney 1997)! On return from Russia to Switzerland, and following his brief trip to Italy with his mentor Carl Vogt (see the preceding section), Penard departed for Wiesbaden, Germany, for another tutorial job, this time with the family of Baron Belevski (years 18871890). Making good use of his time, he did manage to publish, in German, some new observations on rhizopods from the area (e.g., see Penard 1890a,b). In 1891, our busy traveling amoeba-man set out for America, expecting to secure a teaching position at Columbia University, New York City, for which he had hopefully applied. Rejected and dejected, he accepted an invitation to visit the University of Colorado, where the first head of biology there, John Gardiner, was a friend who had chanced to meet him in Geneva some years before. His fascination with the Rocky Mountains and their unique rhizopods, as mentioned in the preceding section of this paper, restored Penards spirits and, after several weeks, he returned to Geneva, although still apparently looking for employment. Next occurred his third tutoring job, his second in Russia. Living with the family of Prince Youssoupoff (years 18921898), he had his hands full as a

teacher of young Felix Youssoupoff, who is especially remembered for his part, later, in plotting the assassination of Rasputin. Penards return to Geneva this time was a doubly happy one: he was departing Russia at a politically tumultuous time in its history and he was bringing back a wife, a lovely Russian woman who had also been employed as a member of Youssoupoffs household. Any other, subsequent visits out of Geneva or Switzerland were of much shorter duration and generally of reduced significance to themes of the present account of Penards life and protistological works.

Later Years and Accumulated Honors

At age 41, Joseph Leidy took a bride without breaking stride. The marriage, coming in 1864, near the end of the Civil War, surprised friends and colleagues who had not even been aware of a courtship in progress with the quiet, plump, motherly woman, daughter of a clergyman from Kentucky. Their marriage was a long and happy one. Childless, Joseph and Anna, in 1876, adopted a charming little girl, Allwina Franck, daughter of a colleague at U. Penn, when her parents both passed away from a long shared illness. A student who, at the time, was living in a room on the third floor of the Leidy house described the trio as always such a charming happy family circle! Leidy continued to work mostly from his home; his amazing powers of concentration let nothing that was going on around him distract him in the least, at times when he thought he required such solitude. Nevertheless, when little Allie once suffered a severe bout of diphtheria, Joseph is said to have sat by her bedside continuously for days and nights, neither sleeping nor eating (Warren 1998). Indefatigable, Leidy in his middle years continued to produce research papers and treatises while taking on more and more outside responsibilities, serving on the boards or directorships of numerous scientific, academic, and philanthropic institutions and organizations in the greater Philadelphia area, actively joining the faculty at nearby Quaker-founded Swarthmore College (in addition to his U. Penn positions), aiding in starting the Philadelphia Zoo (now the oldest in America), etc. He was a man who simply didnt know how to say, No! He even attended social events (if associated with the above activities); he was such an interesting and witty conversationalist (the last man who knew everything: Warren 1998) as well as cutting an imposingly handsome figure in evening wear that everyone, espe-

76

J. O. Corliss

cially the ladies in attendance, always wanted to be seated alongside him at banquet tables. Speaking of women, Leidy contrary to widespread practices in those days (and, alas, even until quite recent years: for the protozoological picture, see Corliss 1993) had no compunctions about having them in his classes in medical school, in institutes where he taught, or in the budding Swarthmore College just outside Philadelphia. Apparently, he was never officially an adviser or mentor, but interestingly enough one of his female students in 1884, Adele M. Field (for 15 years a Baptist missionary in the Orient) was so inspired by his discourses on rhizopod amoebae that she subsequently studied and published papers on these protists from collections made in China. For the most part, Leidys enthusiastic attention to vertebrate fossils during the 1850s, 1860s, and early 1870s eclipsed his precise microscopical researches on protozoological, parasitological, and other materials during those decades, distressing workers around the world in such fields (Ward 1923). The fossil work, however, earned him such titles as the Cuvier of American Paleontology and brought him much (additional) fame from abroad; but it is quite beyond detailed consideration in the present paper (he published nearly 2,000 pages on extinct forms, mostly to be found in these four weighty monographs: Leidy 1854, 1865, 1869, 1873). Perhaps the single most startling early paleontological and evolutionary discovery made by young Leidy (in 1858) and with a correct deduction about its normal posture was that of a nearly complete skeleton from New Jersey of a 28-foot herbivorous duckbilled dinosaur (Hadrosaurus foulkii) deduced by the Philadelphian to have been a biped, the first dinosaur ever so described. When he announced the news, most European paleontologists could/would not believe it, but T. H. Huxley was overjoyed because such a reptile supported his own idea on the origin of birds from dinosaurs. Following appearance of Leidys (1856) unique paper, A Synopsis of Entozoa and Some of Their Ectocongeners (a work containing descriptions of 172 species of parasitic protozoa and worms, many new), it had seemed that he was indeed leaving such a fruitful budding field, parasitology, forever (see the above paragraph). On subsequently returning largely to study of minute living forms, in the late 1870s, it became quite clear that one of the prime reasons for his eventual almost complete withdrawal from the enticing field of vertebrate fossils, sadly enough, was because it had become so unenjoyable, even distasteful, for him in America under its new domination (starting in the 1860s) by fierce

rivals of each other, Marsh of Yale and Cope of Penn (Leidys juniors by 8 and 17 years, respectively, and the latter a student of his), who had became vitriolic in their heated rivalry and competitiveness, were paying large sums for discovered/uncovered bones (which in the past had been saved by farmers and amateur collectors and obligingly sent free to Leidy), were pirating each others collections, rushing into publication even in daily newspapers for priority in describing new forms, etc., etc. So, finally, we come to Leidys mammoth monograph, Fresh-water Rhizopods of North America, published in 1879 by the U. S. Geological Survey. It has become a classic because of the contents of its 324 pages of text material and, perhaps even more, for its 48 large plates of nearly 1,200 exquisitely executed figures in color. In a rare display of pique, the gentle monographer complained that the lithographer hired to prepare his art work for printing didnt do full justice to the original drawings, causing an anxious moment for editor F. V. Hayden. It has been suggested that Leidy was a naturalist-illustrator while the much admired ornithologist Audubon [17851851] was an artist-naturalist, a neat distinction between these two great men with respect to their professional lives and the emphasis shown in their artistic productions (Warren 1998). The still authoritative rhizopod work portrays species of some 40 valid genera (conservatively claimed by Leidy to contain a total of ca. 80 species), while also providing synonyms and other nomenclatural details concerning all forms studied. Modestly, he considered only a relatively few of his described organisms as new to science: later workers, including Penard, tended to split up many of his taxonomically lumped forms. Leidys material was collected from sites along the shores of the Atlantic Ocean and as far west as lands now recognized as the states of Colorado, Nebraska, Wyoming, the Dakotas, and Utah. Gathering material required some trips out of Pennsylvania, although other people also sent collections (i.e., soil samples, etc.) to him from time to time. Cystic as well as trophic stages were depicted. The majority of Leidys descriptions were of species belonging to genera (using names here that are considered correct today) such as Amoeba, Dinamoeba, Endamoeba, and Trichamoeba among the naked amoebae; Arcella, Difflugia, and Euglypha, among test-bearing forms; and Actinophrys, Actinosphaerium, and Clathrulina, among the heliozoa (e.g., see Figures 3 and 4, and the full color reproduction of Leidys Amoeba proteus which adorns the cover of this issue of Protist). Leidys splendid description (in both his text and figures) of the uni-

Two Most Remarkable Amoeba Men

Figure 3. Illustration of species from several sarcodinid genera studied in detail by both Leidy and, a few years later, Penard. All drawings are taken from Leidy (1879), where they appeared in natural colors, a tribute to Leidys artistic abilities. A. Amoeba proteus, perhaps the best-known naked rhizopod amoeba around the world. Its pseudopodia are of the thick lobopodium type. B. Euglypha acanthophora, a thecate or testaceous amoeba with regularly arranged siliceous scales, several spines, and dichotomously branched filopodia. C, D. Also testate (chitinous), Arcella dentata and A. discoides have slender lobopodia; body attached to test by ectoplasmic strands. E. Difflugia amorpha, with test composed of cemented sand grains, sometimes plus diatom shells or other foreign bodies; simple cylindrical lobopodia, occasionally slightly branched. F. Clathrulina elegans, a heliozoan (actinopod sarcodinid) with a brownish envelope perforated by numerous circular or polygonal openings, a long stalk, and numerous fine pseudopodia (of the axopodial type) projecting radially through the pores of the test.

77

78

J. O. Corliss

Two Most Remarkable Amoeba Men

79

versally familiar Amoeba proteus has been widely hailed as the best unambiguous account of this organism; fittingly, Leidy is given the authorship of its name (and as he spelled it), as of 1879. As in the cases of many of his rhizopods, a very brief description of the organism occurred in a note published before appearance of his monograph (in the case of Amoeba proteus, in the year 1878), but subsequent workers have universally prefer to cite 1879 as the date of the origin of many such Leidyan names. Principally because of this supreme work, Leidy has been given still another founder title, the Father of American Protozoology. In 1881, Leidys small but ground-breaking book, The Parasites of Termites, appeared, largely featuring the many symbiotic flagellated protists and bacteria in the gut of the wood-eating hosts. He had always planned, some day, to publish a major treatise on all of the parasites (cestode, trematode, and nematode worms, insects and arachnids, bacteria and protists such as amoebae, flagellates, and ciliates) that he had described briefly and preliminarily over the years, but he never succeeded in carrying out this ambition. Fortunately, his brother Philips son named Joseph Leidy, II and also called Joseph Leidy, Jr. edited a 281-page book published in 1904 by the Smithsonian Institution (Researches in Helminthology and Parasitology) that is essentially a collection of the notes and papers and illustrations (some never published elsewhere) of his uncles works in parasitology: this, to a degree, fulfilled (the first) Joseph Leidys dream in this field of research. Following his stressful last trip to Europe, in 1889, poor Leidy never fully recovered his health. In spite of that, he insisted on maintaining his heavy schedule as director, head, professor, etc.(!) of various institutions and colleges, and on carrying out additional researches which increased his great inventory of Nature (Glassman et al. 1993). In fact, in the short period 1889-1891 (recall that he passed away

on 30 April 1891), he continued to leap hedges, publishing notes and papers on topics as diverse as gregarine protozoa (with many additional observations never completely finished: but see Crawley 1903 for some of Leidys new species), boring sponges, parasitic copepods, ticks, ichneumon flies, avian parasites of the Little Blue Heron, the sabre-tooth tiger and Hippotherium and Rhinoceros from Florida, fossil human bones, and an extinct genus allied to the peccaries. And he brought out the 2nd edition of his textbook on human anatomy at the same time! Also see a posthumous paper (Leidy 1896), which included further data on fossil vertebrates from Florida. Altogether, over some 47 years of active research, our prolific naturalist authored more than 600 publications involving some 2,500 species, a goodly number of which were new to science. Leidys honors were many, and certainly not limited to the inscriptions, statue (designed by Samuel Murray, a student of the renowned painter and sculptor Thomas Eakins), portraits, named streets and laboratories and buildings, etc. to be found in his home town of Philadelphia. But also not to be neglected is mention of the coveted Joseph Leidy Medal, struck in 1923 by the Academy of Natural Sciences of Philadelphia (an organization founded in 1812, eventually publisher of its own journal, and which bestowed lifelong membership on Leidy in 1845) to commemorate the centenary of the birth of their most distinguished member ever. The medal is awarded every three or four years to honor exceptional exploration, research, discovery, and publication in a wide range of scientific disciplines. Its first recipient, the only protistologist ever to receive it to date, was Herbert Spencer Jennings [18681947], famed for his research on behavior and evolution of unicellular organisms (including rhizopod amoebae) and, later, on genetics of the ciliate Paramecium (see Sonneborn 1975). Not surprisingly, Jennings was an admirer of both Leidy and Penard.

Figure 4. More sarcodinids described or redescribed by Leidy and Penard, with figures all from Leidys (1879) colored renditions of the organisms. A. The common freshwater heliozoon Actinosphaerium eichhornii: relatively large body, multinucleate, many radiating axopodia, feeds mostly on diatoms and other algal protists, occasionally a rotifer; ectoplasm and endoplasm both highly vacuolated, with vacuoles larger in the former. B. Trichamoeba villosa, a large naked amoeba characterized by its one to several broad lobopodia and a semi-permanent uroid with posteriorly projecting fine filaments or villi; clear ectoplasm, one nucleus, one contractile vacuole, several food vacuoles. C. Baileya mutabilis, a thecate rhizopod with a clear and flexible test and long filopodia that may branch, cross, and/or even anastomose. Single contractile vacuole. D. Dinamoeba mirabilis, a naked amoeba with numerous long, clear, conical/tapering lobopodia that twist or coil in retraction. Entire body surface often bristles with a covering of attached bacteria. (As is also true of figures AF of my preceding Figure 3; figures AD here have been retouched slightly to aliminate disconcerting pseudopodia coming from other (neighboring) figures on Leidys original plates that have invaded the space occupied principally by the organism specifically being portrayed).

80

J. O. Corliss

Leidy received an honorary doctorate from Franklin and Marshall College in 1869, and from Harvard University in 1886. In 1880, the Boston Society of Natural History bestowed the annual Walker Grand Prize on him. He was among the 50 leading men of science to establish the National Academy of Sciences in 1863. From abroad, in addition to invited membership in various long-distinguished societies, he was elected to honorary membership in the Royal Microscopical Society of London in 1879 and in the Zoological Society of London in 1883. In 1884, the Geological Society of London awarded him its Lyell Medal; and, two years later, the Institute of France bestowed its Cuvier Prize Medal on him. Finally, mention should be made of taxonomic names, patronyms, created in his honor. For example, some gregarine sporozoa from crickets have been placed in the genus Leidyana Watson, 1915, a group now with its own familial name, Leidyanidae Kudo, 1954; and Leidyella Jung, 1942, was proposed for a genus of testaceous rhizopod amoebae, but has been little used since. The generic names Leidyonella and Leidyopsis were at one time suggested for one of Leidys species of a parasitic flagellate in termites, but Kudo (1966) considers them to be junior synonyms of Trichonympha Leidy, 1877. Although I have decried the fact that Joseph Leidy seems little remembered personally today, there were many eulogies delivered internationally around the time of his passing (year 1891), and the great paleontologist Henry Fairfield Osborn eventually produced a glowing report in his biographical memoir for the National Academy of Sciences (Osborn 1913). On the centenary of his birth, W. S. Middleton, M.D. (1923) praised him mightily in the Annals of Medical History, as did H. B. Ward, Ph.D. (1923) in the Journal of Parasitology (a journal founded by Ward who, himself, has often been hailed as the Father of American Parasitology, which would make Leidy the Grandfather?). And recall the book of Leidys parasitological notes edited by his nephew (Leidy, Jr. 1904). On/near the centenary of Leidys death, Susan Glassman et al. (1993) published a fine honorific work, in Philadelphias own Proceedings of the Academy of Natural Sciences, including an Appendix on the number of genera and species he worked with from each of the major taxa of organisms. D. H. Wenrich, parasitologist and protozoologist, an illustrious 20th century successor of Leidys in Zoology at the University of Pennsylvania, had amassed copious data on his celebrated predecessor with the intention of publishing a lengthy biography; alas, his plans were interrupted by his own sud-

den passing, in 1968, at the age of 83. Finally, Leonard Warren, M.D., Ph.D. (1998), Professor at both the Wistar Institute and (emeritus) U. Penn, has produced the first long-awaited full-length biography (303 pp.) of Leidy, truly a sensitive and understanding masterpiece. At age 43, Eugne Penard brought back a bride from Russia. It is interesting to note that his age was almost the same as that of Leidys when the latter took the same matrimonial plunge. M. et Mme Penard remained a happy couple until her passing, at age 89, in 1945: 47 years together compared with Leidys 27 years of marital bliss, since the American rhizopodologist died at the relatively tender age of 68. Back to another parallel: the Penards apparently had no children of their own. Penards active protistological research years, 18881922, quantitatively about matched Leidys non-paleontological total because, although the Genevan still had the last third of his (considerably longer) life to go, his eyesight fatigue brought practically to a halt all detailed microscopical work beyond 1922. Penards second major monograph on free-living amoebae, after the earlier one of 1890 cited in a preceding section of this paper, came out nine years later (Penard 1899); the depths of Lake Geneva served as the collecting site this time. It was quite soon followed with several shorter important works in between by his huge (714 pages!) Faune rhizopodique du bassin du Lman (Penard 1902), a most authoritative taxonomic-ecological survey with many new species described in greater detail than that ever seen in most older works, including Leidys. In a complimentary way, the Swiss protistologist might be referred to as a master of minutiae. Penards first monograph on the freshwater hliozoaires appeared in 1904, quickly followed by two on the sarcodins sensu lato (Penard 1905a,b). During the same and the next seven years, a considerable number of additional substantial works (some on edaphic and terrestrial as well as aquatic forms) appeared, including observations on material collected (or sent to him) from such sites as neighboring countries, the British Isles (including the celebrated Loch Ness, home of the alleged monster, in Scotland), Colombia (via a German expedition to South America), the Himalayan mountains of India, and South Pole regions (collections of materials gathered during early Antarctic expeditions carried out by both British and French explorers). Such a fantastically busy microscopist was this humble rhizopodologist! In 1914, in the 60th year of his life, the seemingly tireless ( la Leidy!) Penard commenced his prolific

Two Most Remarkable Amoeba Men

81

series of investigations on predominantly freshwater free-living (free-swimming or attached to some living or inanimate substrate) ciliated protozoa, including the suctorians (then considered a high-level separate taxonomic group, the Tentaculifera: see Corliss 1979). The year had actually started, however, with an account on the rotifers of Switzerland, including a description of several new species of these tiny micrometazoan organisms. The protozoan researches essentially culminated in a monographic paper on the suctorians (Penard 1920) and a monographic book on the regular ciliates (Penard 1922a); the latter included a detailed treatment of some 300 species of which 168 were new. In between these authoritative publications, Penard (1921) managed to complete a paper on flagellates which appeared (by invitation, perhaps?) in the Proceedings of the Academy of Natural Sciences of Philadelphia, a favorite outlet of Leidys, as the reader will recall. By the end of the year 1922, Penard had published some 2,150 pages (exclusive of plates of figures) on the protozoa (primarily sarcodinids) in his eight major monographs alone! His total number of scientific publications, spanning his less than 30 years of active research on microorganisms, comes to ca. 78 (eight each in German and English, the remainder in French). The total number of protist species he described as new amounts to ca. 530, with slightly over 62% being sarcodinids (Deflandre 1958). Like Leidy, Penard worked mostly at the species level; neither of them described very many new genera, usually content to stretch the boundaries of the well known genera of such authoritative predecessors as Ehrenberg, Dujardin, and Hertwig. As mentioned earlier, the medically enforced retirement as a microscopist obliged Penard at age 67 but more than 30 years from the eventual date of his demise to give up his passionate activity as an astute observer of living protists. Presumably, he took this pronouncement philosophically and goodnaturedly. He turned to writing some review articles and some more (the first ones had appeared in 1905) adventure and natural history novels and books for young people (for the last one, see Penard 1946); and he re-arranged his many boxes of slides of type-material of the numerous new species (see above) he had described. These slides, mostly of carmine-stained sarcodinids and ciliates, number well over one thousand and are to be found today distributed among three major museums of natural history, all in Europe: the ones in London, Paris, and Geneva. In 1952, I examined some of these myself (as had Georges Deflandre before me: see Deflandre 1954, 1958) and was amazed at their clarity despite their ages of 30-50 years or more!

Penard (19401941) also entered the realm of animal psychology, attributing behavioral traits to protists: for example, instinct and memory. Some of such investigations/thoughts on these possible characteristics of protozoa are perhaps best succinctly summarized in one of the suppositions proposed in his last paper, written in English and entitled Habituation (Penard 1947), which may be paraphrased as follows: the protozoa know what they are doing! One of his examples: loricate ciliates (under microscopical examination) will quickly withdraw/contract into their tests when the coverslip is lightly tapped; but that response lessens on repeated tappings. This paper was preceded by two books by Penard (1922b, 1938) setting the stage for his insightful entrance into the vast field of behavior of lower organisms (recall Jennings 1904, 1931), which Penard felt was unfairly neglected by the great British psychologist W. H. Thorpe. Penard had met Jennings (his junior by some 13 years) once in Europe, around the turn of the century. In 1945, following the death of his wife, Penard at the age of 89 moved into a rest home (principally for elderly ladies, but he was deemed harmless!?) on the outskirts of Geneva. He remained intellectually sharp, multilingual, and steady in voice and writing hand, his weakened visual acuity notwithstanding. Nine years later, on 5 January 1954, he quietly passed away. Recognition of Eugne Penards invaluable contributions to protozoology came to him at various times in his long life. The Faculty of Sciences of Geneva twice bestowed awards on him (in years 1922, 1945), and he was elected to membership in leading biological (including the Socit suisse de Microbiologie) and natural history societies in his home country. He held the office of President of Honor of the Socit suisse de Zoologie from 1936 until his death. Though officially an amateur biologist, he was accorded the fame and respect usually reserved for professionals (the category to which Joseph Leidy and the great majority of renowned scientists over the ages have belonged). Internationally, Penard was widely recognized as well, principally for his abilities in microscopy in the study of living protozoological materials. He was made a (foreign) Fellow of the Linnean Society of London and of the Royal Microscopical Society of London, and a member of the renowned Queckett Club in London. He was early elected a Membre dHonneur de la Socit franaise de Microscopie. In 1952, a scant two years before his death, he became the first Honorary Member of the young Society of Protozoologists, headquartered in America but international in membership (Corliss 1998). This

82

J. O. Corliss

last-mentioned honor pleased him mightily (Corliss 1954). He penned a letter to me including these touching words, This announcement was for me an immense satisfaction for such an honorable proof of what I might have done in protozoan studies And it came from America! Recall that 60 years earlier Penard had spent several enjoyable months collecting amoebae in the Rocky Mountains of the United States, his only trip to America. Half a dozen generic names that honor the modest Swiss protistologist have been given to protozoa, five to rhizopod amoebae and one to a ciliate. In chronological order, they are Penardia Cash, 1904, Penardiella Jung, 1942, Penardochlamys Deflandre, 1953, Penardogromia Deflandre, 1953, Penardeugenia Deflandre, 1953, and Epenardia Corliss, 1971. I like to think that the last, a replacement name for a ciliate first seen and well described by Penard as a Glaucoma species always found in the vicinity of the aquatic plant Myriophyllum, would have evinced an amused smile from the good humored Swissman because of the play on words in its name: epinard is the French word for spinach. To my knowledge, Penard as a person has received little attention of a biographical nature in the literature since his passing in 1954. Aside from the short necrological notices appearing at the time in several European journals, only the rather brief papers of Deflandre (1954, 1958) and Corliss (1954, 1956) mention aspects of his personal life while also emphasizing his importance to protozoology.

cludes mention of 10 of Leidys genera and more than two dozen of his species, and 15 of Penards genera and nearly four dozen of his species, descriptions all originally made scores of years earlier and still holding up today. Specialists are familiar with many more. Numerous books and compendia reproduce figures originally published by Leidy and Penard. Thus, one might conclude that their works are nearly as immortal as the protists they studied with such earnest ardor and open admiration over 100 years ago.

References
N.B. Included below, among the rest of the literature citations, are only some 14 works by Leidy and two dozen by Penard, out of their numerous combined total. Most of these are directly cited (in the customary/conventional manner) on preceding text-pages, while a few are just indirectly referred to in the body of this essay but included here with the others for the readers interest. Not surprisingly, many of Leidys papers, including major ones, are clearly of a non-protistological nature (see text). Data on numbers of pages and figures are sometimes purposely supplied, especially for Leidy whose works are often known for their illustrations. Awerinzew S (1906) [Freshwater Rhizopoda] II. Trav. Soc. Nat. St. Ptersbourg 36 (2): 121346 (in Russian) Bary A de (1859) Die Mycetozoen. Z Wiss Zool 10: 88175

Concluding Comments
Both Joseph Leidy of Philadelphia and Eugne Penard of Geneva were exemplary men in their display of unbounded energy, incredible powers of concentration, high visual acuity, love for and amazing memory of details, and unrestricted passion in pursuit of the lowly amoebae. Above all, perhaps, they were humble non-fame-seeking naturalists who, without the advantages of modern technology, turned their enthusiastic attention to the full bionomics (taxonomy plus ecology) of the rhizopod sarcodinids to the lasting benefit of protozoology. Their personal lives, interesting though they may have been, are not as important to posterity, of course, as their scientific contributions to the disciplines or fields within which they worked. Many of their observations adding to the great inventory of Nature are not likely to be forgotten: consider the fact that such distinguished protistologists as R. R. Kudo [18861967], in the last edition of his widely used textbook of protozoology (Kudo 1966), in-

Blochmann F (1886) Die Mikroskopische Pflanzen- und Thierwelt des Ssswassers. Braunschweig Bovee EC, Jahn TL (1973) Taxonomy and phylogeny. In Jeon KW (ed) The Biology of Amoeba. Academic Press, New York and London, pp. 3782 Btschli O (18801882) Protozoa. Abt. I. Sarkodina und Sporozoa. In Bronn HG (ed) Klassen und Ordnung des Their-Reichs, CF Winter, Leipzig, 1: 1616 Cash J, Hopkinson J (1905) The British Freshwater Rhizopoda and Heliozoa. Vol. 1, Part I. Ray Society, London Cash J, Hopkinson J (1909) The British Freshwater Rhizopoda and Heliozoa. Vol. 2, Part II. Ray Society, London Cash J, Wailes, GH (1919) The British Freshwater Rhizopoda and Heliozoa. Vol.4. Ray Society, London Cash J, Wailes GH (1921) The British Freshwater Rhizopoda and Heliozoa. Vol. 5. Ray Society, London Cash J, Wailes GH, Hopkinson J (1915) The British Freshwater Rhizopoda and Heliozoa. Vol. 3. Ray Society, London

Two Most Remarkable Amoeba Men

83

Cienkowski L (1863) Das Plasmodium. Jahrb Wiss Bot 3: 400441 Claparde E, Lachmann J (18581861) Etudes sur les Infusoires et les Rhizopodes. Mm. Inst. Nat. Genvois. [Sole section on les rhizopodes is in Mmoire No. 6, pp 413466] Cole FJ (1926) The History of Protozoology. University of London Press, London Corliss JO (1954) Dr. Penard and America. J. Protozool. 1: 191 Corliss JO (1956) Dr. Eugne Penard. Obituary. Proc Linn Soc London, pp 4244 Corliss JO (1975) Three centuries of protozoology: a brief tribute to its founding father, A. van Leeuwenhoek of Delft. J Protozool 22: 37 Corliss JO (19781979) A salute to fifty-four great microscopists of the past: a pictorial footnote to the history of protozoology. Parts I and II. Trans Amer Microsc Soc 97: 419458; 98: 2658 Corliss JO (1979) The Ciliated Protozoa: Characterization, Clasification, and Guide to the Literature. 2nd ed. Pergamon Press, London and New York Corliss JO (1992) Historically important events, discoveries, and works in protozoology from the mid-17th to the mid-20th century. Rev Soc Mex Hist Nat 42 (year 1991): 4581 Corliss JO (1993) The contributions of women to the science of protozoology. Acta Protozool 32: 129134 Corliss JO (1998) The golden anniversary of the Society of Protozoologists (19471997). J Euk Microbiol 45: 126 Crawley H (1903) The polycystid gregarines of the United States. Proc Acad Nat Sci Phila 55: 4158, 632644 Deflandre G (1953) Ordres des Testacealobosa (De Saedeleer, 1934), Testaceafilosa (De Saedeleer, 1934), Thalamia (Haeckel, 1862) ou Thcamoebiens (Auct.) (Rhizopoda Testacea). In Grass P-P (ed) Trait de Zoologie, Masson & Cie, Paris Vol 1, fasc 2, pp 97148 Deflandre G (1954) Eugne Penard (18551954), sa vie et son oeuvre. J Protozool 1: 187190 Deflandre G (1958) Eugne Penard (18551954) Correspondance et souvenirs. Bibliographie et bilan systmatique de son oeuvre. Hydrobiologia 10: 137 Dobell C (1919) The Amoebae Living in Man: A Zoological Monograph. J Bale, Sons & Danielsson, London Dujardin F (1841) Histoire des Zoophytes. Infusoires. Paris Ehrenberg CG (1838) Die Infusionsthierchen als Vollkommene Organismen. Leipzig Glassman S, Bolt Jr EA, Spamer EE (1993) Joseph Leidy and the Great Inventory of Nature. Proc Acad Nat Sci Phila 144: 119

Haeckel E (1878) Das Protistenreich Gnther, Leipzig Haeckel E (1894) Systematische Phylogenie I. Systematische Phylogenie der Protisten und Pflanzen. G Reimer, Berlin Hertwig R (1879) Organismus der Radiolarien. Denkschr Med-Naturwiss Gesellsch 2: 1227 Hertwig R, Lesser E (1874) ber Rhizopoden und denselben nahestehende Organismen. Arch Micros Anat 10 (Suppl): 35243 Jennings HS (1904) Contributions to the study of the behavior of lower organisms. Carnegie Inst Wash Publ No. 16: 1256 Jennings HS (1929) Genetics of the Protozoa. Bibliogr Genet 5: 105330 Jennings HS (1931) Behavior of the Lower Organisms. Rev ed. Columbia University Press, New York Joblot L (1718) Descriptions et Usages de plusieurs nouveaux Microscopes avec de nouvelles Observations. Paris Kudo RR (1966) Protozoology. 5th ed. Charles C Thomas, Springfield, IL Lee JJ, Hutner SH, Bovee EC (eds) (1985) An Illustrated Guide to the Protozoa. Society of Protozoologists, Lawrence, KS Lee JJ, Leedale GF, Bradbury PC (eds) (2001) An Illustrated Guide to the Protozoa. 2nd ed. Society of Protozoologists, Lawrence, KS Leidy J (1844) On the comparative Anatomy of the Eye in Vertebrated Animals [Thesis for M.D. degree, University of Pennsylvania Medical School] Leidy J (1853) A Flora and Fauna within Living Animals. Smithsonian Contrib Knowl 5, 68 pp, 10 pls Leidy J (1854) The ancient Fauna of Nebraska, a Description of extinct Mammalia and Chelonia from the Mauvaises Terres of Nebraska. Smithsonian Contrib Knowl 6, 126 pp, 25 pls Leidy J (1856) A synopsis of Entozoa and some of their ecto-congeners, observed by the author. Proc Acad Nat Sci Phila 8: 4258 Leidy J (1858) Hadrosaurus foulkii, a huge extinct herbivorous saurian. Proc Acad Nat Sci Phila 10: 215218; reprinted in Amer J Sci 26 (2nd ser.): 266270 Leidy J (1861) An Elementary Treatise on Human Anatomy. 1st ed. J B Lippincott, Philadelphia Leidy J (1865) Cretaceous reptiles of the United States. Smithsonian Contrib Knowl 14, 140 pp, 20 pls Leidy J (1869) The extinct mammalian fauna of Dakota and Nebraska, including an account of some allied forms from other localities, together with a synopsis of the mammalian remains of North America. J Acad Nat Sci Phila 7: 1472, plus 39 pls Leidy J (1873) Contributions to the extinct vertebrate fauna of the Western Territories. Rep U.S. Geol Surv Terr 1, 358 pp, 37 pls

84

J. O. Corliss

Leidy J (1879) Fresh-water Rhizopods of North America. U.S. Geol Surv Terr Rep 12: 1324, plus 48 pls in color Leidy J (1881) The Parasites of Termites. Collins, Philadelphia Leidy J (1889) Treatise on Human Anatomy. 2nd ed. J B Lippincott, Philadelphia Leidy J (1896) Fossil vertebrates from the Alachua clays of Florida Mastodon, Rhinoceros, Hippotherium, Procamelus. Trans Wagner Free Inst Sci, year 1896, 47 pp, 19 pls. [a posthumous publication edited by F A Lucas five years after Leidys death] Leidy Jr J (ed) (1904) Researches in Helminthology and Parasitology. Smithsonian Misc Coll 46: 1281 [theretofore unpublished notes by Joseph Leidy, his uncle, on diverse parasitological materials] Lister A & G (1925) Monograph of the Mycetozoa. 3rd ed. Brit Mus Nat Hist, London Margulis L, Corliss JO, Melkonian M, Chapman DJ (eds) (1990) Handbook of Protoctista. Jones and Bartlett, Boston Maupas E (1888) Recherches exprimentale sur la multiplication des infusoires cilis. Arch Zool Exp Gn (sr. 2) 1: 165277 Maupas E (1889) Le rajeunissement karyogamique chez les cilis. Arch Zool Exp Gn (sr. 2) 7: 149517 Middleton WS (1923) Joseph Leidy, scientist. Ann Med Hist 5: 100112 Mller J (1858) ber die Thalassicollen, Polycystinen und Acanthometren des Mittelmeeres. Abh Akad Wiss Berlin 43: 162 Mller OF (1786) Animalcula Infusoria Fluviatilia et Marina. Havniae et Lipsiae Nanney DL (1997) Frontier connections. Rev Soc Mex Hist Nat 47: 201216 Osborn HF (1913) Biographical memoir of Joseph Leidy, 18231891. Biog Mem Nat Acad Sci USA 7: 335396 Penard E (1888) Recherches sur le Ceratium macroceros [Thesis for D.Sc. degree, University of Geneva] Penard E (1889)a Etudes sur quelques Hliozoaires deau douce. Arch Biol 9: 419472 Penard E (1889)b Notes sur quelques Hliozoaires. Arch Sci Phys Nat Genve 22: 523539 Penard E (1890)a ber einige neue oder wenig bekannte Protozoen. Jahrb Nassau Verh Naturkd 43: 7591 Penard E (1890)b Die Heliozoen der Umgegend von Wiesbaden. Jahrb Nassau Verh Naturkd 43: 3966 Penard E (1890)c Etudes sur les Rhizopodes deau douce. Mm Soc Phys Hist Nat Genve 31: 1230 Penard E (1891)a Les Pridiniace du Lman. Bull Trav Soc Bot Genve, year 1891, pp 163

Penard E (1891)b Contributions ltude des Rhizopodes du Lman. Arch Sci Phys Nat Genve 26: 134156 Penard E (1891)c Rocky Mountain rhizopods. Amer Nat 25: 10701083 Penard E (1893) Pelomyxa palustris et quelques autres organismes infrieurs. Arch Sci Phys Nat Genve 29: 165182 Penard E (1899) Les Rhizopodes de faune profonde dans le lac Lman. Rev suisse Zool 7: 1142 Penard E (1902) Faune rhizopodique du bassin du Lman. Kundig, Genve [714 pp] Penard E (1904) Les Hliozoaires deau douce. Kundig, Genve [341 pp] Penard E (1905)a Les Sarcodins. In: Catalogue des Invertbrs de la Suisse. Mus Hist Nat Genve 1: 1164 Penard E (1905)b Les Sarcodins des grands lacs. Kundig, Genve [136 pp] Penard E (1914)a A propos de Rotifres. Rev suisse Zool 22: 125 Penard E (1914)b Les Cothurnids muscicoles. Mm Soc Phys Hist Nat Genve 38: 1965 Penard E (1914)c Sur quelques Tentaculifres muscicoles. Arch Protistenkd 34: 277294 Penard E (1920) Etudes sur les Infusoires Tentaculifres. Mm Soc Phys Hist Nat Genve 39: 131229 Penard E (1921) Studies on some Flagellata. Proc Acad Nat Sci Phila 73: 105168 Penard E (1922a) Etudes sur les Infusoires deau douce. Georg & Cie, Genve [331 pp] Penard E (1922b) Les Protozoaires considrs sous le rapport de leur organique. Georg & Cie, Genve [95 pp] Penard E (1938) Les infiniment petits dans leurs manifestations vitale. Georg & Cie, Genve [212 pp] Penard E (19401941) Protozoaires et psychologie. Arch Sci Phys Nat Genve 22: 160175, 179200, 203226, 265289; 23: 5776 Penard E (1946) La Roche aux Echidns. Delachaux et Niestl, Neuchatel and Paris [Penards last book for children, illustrated by his own line drawings] Penard E (1947) Habituation. J Roy Microsc Soc 67: 4345 [in English] Perty M (1852) Zur Kenntniss kleinster Lebensformen nach Bau, Functionem, Systematik, mit Specialverzeichniss der in der Schweiz, beobachteten. Jent & Reinert, Bern Rhumbler L (1903) Systematische Zusammenstellung der recenten Reticulosa (Nuda + Foraminifera). I. Theil. Arch Protistenkd 3: 181294 Ride WDL (1999) Introduction to The International Code of Zoological Nomencature, 4th ed. International Trust for Zoological Nomenclature, London, pp. xixxxix

Two Most Remarkable Amoeba Men

85

Rsel von Rosenhof AJ (1755) Der Monatlich-Herausgegebenen Insecten-Belustigung. 3 (Suppl.). Nrnberg Schaeffer AA (1916) Notes on the specific and other characters of Amoeba proteus Pallas (Leidy), A. discoides spec. nov. and A. dubia spec. nov. Arch Protistendk 37: 204228 Schaeffer AA (1920) Ameboid Movement. Princeton University Press, Princeton, NJ Schaeffer AA (1926) Taxonomy of the Amebas. Papers Dept Mar Biol Carnegie Inst Wash 24: 1116 Schewiakoff W (1926) Acantharia. Fauna e Flora del Golfo di Napoli. Staz Zool Napoli, Monogr No. 37: 1775 (in German)

Schulze E (1874) Rhizopodienstudien. Arch Mikros Anat 10: 1328 Sonneborn TM (1975) Herbert Spencer Jennings. Biog Mem Nat Acad Sci USA 47: 143223 Wailes GH (1912) Freshwater Rhizopoda and Heliozoa from the States of New York, New Jersey and Georgia, U.S.A. J Linn Soc Zool 32: 121161 Wailes GH (1913) Freshwater Rhizopoda from North and South America. J Linn Soc Zool 32: 201218 Ward HB (1923) The founder of American parasitology, Joseph Leidy. J Parasit 10: 126 Warren L (1998) Joseph Leidy, the Last Man Who Knew Everything. Yale University Press, New Haven and London