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A • G. Evelyn Hutchinson, Yale University • C 1954 l e n i a t e n n

The progress of Man in civilization, no less than his numerical increase, continually extends the geographical domain of Art by trenching on the territories of Nature, and hence the Zoologist or Botanist of future ages will have a much narrower field for his researches than that which we enjoy at present. It is, therefore, the duty of the natu- ralist to preserve to the stores of Science the knowledge of these extinct or expiring organisms, when he is unable to preserve their lives; so that our acquaintance with the marvels of Animal and Vegetable existence may suffer no detriment by the losses which the organic creation seems destined to sustain.

hus wrote Strickland in the there is a suspicion that before its de- Tintroduction to his classical mise it adopted some of the manners memoir [1] on the dodo, published in of the court of Louis XIV. The birds 1848. It is both extraordinary and tragic from Reunion, without bones to rep- how little we still know about animals resent them, are solely known from the which have become far rarer than they reports of a few travelers and from a were at the time Strickland wrote. No number of problematic pictures and satisfactory treatment of L’éléphant et ses may, in part, be fabulous. amours, let alone a stately ten-volume The existing information about these Grundriss der Elefantenlehre appear, in strange creatures has recently been spite of the popular story, yet to exist. summarized by the late Viscount Ha- Of the rhinoceroses, of which perhaps chisuka, who died a few days before his not more than a few thousand speci- work appeared [2]. His book provides mens of the commonest species are an admirable excuse for an excursion now living, we know even less. The lit- into one of the oddest regions of natural tle that we do know is, however, abun- history. dantly worth knowing. Of the dodo we The three islands which were the cannot now learn much that is new; homes of the birds under consideration what is ascertainable will be discussed lie west of southern Madagascar, as in the following pages. shown in Figure 1. They were probably known to the Arabs but were first The Dodo and the Solitaire sighted by European navigators early No animals that have ever lived seem in the sixteenth century. All were well to have balanced more precariously on forested, lacked land mammals other the boundaries of the real and the imag- than bats, and supported a number of inary than the flightless birds which birds, mostly now extinct, as well as are placed scientifically in the family giant tortoises. Of the last named, three Raphidae. Now, unfortunately, no one species are supposed to have lived on Editors’ Note: The renowned ecologist G. Evelyn can observe their behavior. They lived Rodriguez and a considerable number, Hutchinson was the first scholar to pen “Margi- on the islands of the Mascarene group, perhaps not all contemporary, have nalia” in American Scientist, beginning in 1942. Reunion, Mauritius, and Rodriguez. been described from subfossil or fossil This entry is from his last year as Marginalist, The dodo of Mauritius, as everybody material from Mauritius. Hogs, goats, 1954. The controversy he describes has only been knows, has also been an inhabitant of and chickens were liberated on Mau- resolved within the past decade, and we are not about to spoil the punch line here. If, however, you Wonderland since 1865, the very year ritius in the sixteenth century, but the visit the American Scientist website—after read- in which the best skeletal remains of island remained uninhabited until 1638 ing the Yale professor’s analysis, please—you can the bird were discovered and sent to when the Dutch established a settle- find a selection of links to material about the inevi- Europe. The solitaire never successful- ment. Reunion was apparently visited table reach of genetic analysis to solve the dilemma. ly left its home in Rodriguez, though for timber early in the seventeenth cen-

158 American Scientist, Volume 100 © 2012 Sigma Xi, The Scientific Research Society. Reproduction with permission only. Contact [email protected]. a curiously gaunt and perhaps feather- less bird. Oudemans supposes that this appearance was assumed seasonally, presumably after breeding. Hachisuka thinks that part of the sheath of the beak was also shed at the moult. The contemporary accounts indicate that the bird lived in forests, fed on fallen fruits of some sort, and laid one white egg about 10 cm long in a nest on the ground. The raphid bird of Rodriguez, known generally today as the Solitaire, or scientifically Pezophaps solitaria (Gmelin), is in some ways the best known of the group, because it was the only species that came under the eye of anyone who was really interested in observing it. Un- fortunately, this gentleman, M. François Leguat, who went to Rodriguez in 1691 at the head of the Huguenot settlers, was by his own admission an inexpert draughtsman and so only published one detailed delineation of the bird from life. This is the whole of the icono- graphic evidence for its appearance, but the general form and some details of his figure have been confirmed by a great quantity of skeletal material recovered during the last century. There is, however, a special complication to be faced in considering the solitaire, not present in the case of the birds of the other islands. It has been seriously proposed that, unlike his bird, Leguat himself is a mythical creation. In the Voyage et avantures de François Leguat & de ses Compagnons en deux isles désertes des Indes Orientales [3] the solitaire is described as taller than a tur- key with similar feet and beak, though the latter was a little more hooked. The male is described as grayish and brown, Fig. 1. Redrawn from contemporary sources by Martha M. Dimock [6]. Reproduced by the female as either blonde or brown; Strickland and Hachisuka. one may perhaps suspect an age dif- ference here. The wings were too small for flight; in the account of the male it is said that there was a bony mass tury and was inhabited sporadically skillful and detailed oil paintings and under the feathers as big as a musket after 1649. Rodriguez was supposedly provide a great deal of information as to ball, used in defense. The female had first colonized in 1691 by a small group the appearance of the bird. About a doz- “une espèce de bandeau comme un of Huguenots who attempted to estab- en complete skeletons and many iso- bandeau de veuves en haut du bec qui lish a settlement. lated bones have been recovered from est de couleur tanée.” It is thus clear Though no skin of the Dodo still Mauritius in the past hundred years. from the original, though not from con- exists—Oxford University having de- Most of the illustrations indicate an ex- temporary translations, that the beak stroyed the only specimen as too moth tremely clumsy, fat, usually dark gray or was tan-colored. The band-like struc- eaten in 1755—the Mauritius species, blackish bird with a huge beak, minute ture is clearly dark and conspicuous Raphus cucullatus Linnaeus, is very well functionless wings, and short plumelike in Leguat’s figure. The most striking known. About twenty-five illustrations tail. It has been supposed that the fatter feature of the female, however, was that made during the seventeenth century specimens with less conspicuous tails, at the base of the neck there were two are in existence; these seem mostly to darker plumage, and brown rather than elevations, with whiter feathers than have been executed from captive birds golden irides are females, though no the rest, “qui représente merveilleuse- brought to Europe. Some of these pic- contemporary observer noted any sex- ment un beau sein de femme.” These tures, notably by Roelandt Savory, are ual dimorphism. Some drawings show structures, which must have put Leguat www.americanscientist.org © 2012 Sigma Xi, The Scientific Research Society. Reproduction 2012 March–April 159 with permission only. Contact [email protected]. in mind of some of the ladies he had lawyers, or priests. This observation whole of Leguat’s work, claiming that seen dressed in the fashions of late sev- of the marriage is apparently unique it was a novel written by F. M. Mis- enteenth century France, evidently con- in avian ethology; Leguat’s credibility son. In Atkinson’s attack on Leguat, tributed to his idea that “La femelle est is therefore a matter of considerable the unhappy solitaire was dealt many d’une beauté admirable,” though the interest. undeserved blows. blonde plumage was doubtless most at- Leguat’s book has had a most curi- Atkinson claimed that “Leguat,” that tractive. Of such birds walking to meet ous history. Written in French, it was is, Misson, had invented the solitaire, him with so proud a display he writes first published in England in 1708. A using as his sources the two brief de- “on ne peut s’empêcher de les admirer number of editions in French, English, scriptions by Carré and DuBois of a & de les aimer, de sorte que souvent bird of the same name in Reunion, a de- leur bonne mine leur a sauvé la vie.” scription of the “oiseau de Nazareth,” This statement from the first settler on apparently a dodo, seen on Mauritius an uninhabited island, in the presence The single egg and the by Cauche, other notes on Mascarene of an edible bird—”le goût en est excel- birds given by Cauche and DuQuesne, lent sur tout quand ils sont jeunes”—is long period required and finally, for the peculiar skeletal fea- a tribute to the elegance of the bird and tures of the wing, some experience with the sensitivity of the observer. to rear the young are the skeletons of swans and other Euro- Leguat continues with an account of pean species. The remark, for instance, the territorial behavior of these birds. reasonable in view about the bandeau on the tan-colored The breeding territory had a radius beak is supposedly derived from Cau- of about two hundred yards, and was of the anatomical che—”Leur perdris ont le bec rouge, il y defended during incubation and the en a de tannées.” The practiced philolo- long period of several months while evidence that these gist will doubtless see the inevitability the young bird was helpless. Leguat of the comparison; the present writer says that males drove off intruding birds were derived does not. To put the solitaire together in males and females intruding females; a convincing and, in part, osteologically this is reasonable, but it is less likely from pigeons. confirmed form, would be a precog- that the male called the female when nitive feat placing Leguat as the most he saw an intruding female and vice successful imaginative palaeontologist versa, as Leguat believed that he had who has ever existed. Since Aktinson observed several times. The wing is wrote, the researches of Vivielle and said to have been rotated, making a and Dutch followed in the same year Dehérain have shown that there is in- noise like a rattle when the birds were and the work evidently caused consid- dependent evidence for the reality of calling. This was presumably a territo- erable excitement. Right from the begin- Leguat’s voyage in the maritime ar- rial warning. The nest consisted of a ning doubts as to the authenticity of the chives of Texel, Cape Town, and Mau- heap of palm leaves; a single egg, big- work were raised, and though an obitu- ritius and in some contemporary Dutch ger than that of a goose, was laid; both ary notice exists, no other independent naval correspondence; Mortensen has sexes incubated. Except for what is evidence of Leguat’s reality was known shown that some of Aktinson’s geo- said about the one sex calling its mate until fairly recently. The book was for graphical and biological criticisms are to drive off an intruder of the opposite long the only important source for the invalid. Nevertheless, in spite of all this, sex, this is all reasonable, though in existence and nature of the Rodriguez there is still a tendency to cast doubt on Leguat’s time it would have seemed solitaire and was accepted as such by Leguat’s testimony. Hachisuka’s book less reasonable than it does to us today. Strickland, though by 1848 bones of reviews, if a little unsystematically, the Leguat is clearly one of the founders the solitaire had already been discov- whole problem, and comes out strongly of our knowledge of territoriality in ered. When later complete skeletons in Leguat’s favor. The territorial behav- animals. The single egg and the long of Pezophaps became known, Leguat’s ior recorded, which Atkinson dismisses period required to rear the young are remarks about the “petite masse ronde most unceremoniously, seems to the reasonable in view of the anatomical comme une balle de mousquet” under present writer greatly to strengthen the evidence that these birds were derived the wing feathers received confirma- case for Leguat’s reliability, since there from pigeons. tion. In view of this skeletal verification, were practically no sources available as His description of the end of the Alfred Newton and others felt that the hints of such behavior in 1708. Perhaps rearing of the young, and of the for- work had been vindicated and a reprint the marriage of the solitaires also really mation of mated pairs, which he says was published by the Hakluyt Society occurred. were stable during life, is much odder. in 1891. Practically nothing has been The identity of the bird on Reunion Some days after the young left the nest, added to our knowledge of the soli- raises a problem which has still not a company of thirty or forty brings an- taire since then, though the osteological been solved. There are four contem- other young bird to it, and, the parents evidence on the one hand and Leguat’s porary descriptions. Tatton, who was joining the band, the two young birds account on the other have been usually in Reunion in 1613, writes of “a great are led off to an unoccupied territory considered as providing a fairly good fowl of the bigness of a Turkie, very and left there by the adults. He refers understanding of the bird. fat, and so short winged that they can- to this as a “marriage,” and goes on to In a series of works in the nineteen not flie, beeing white and in a manner moralize on the simple fidelity of these -wenties an American scholar, G. At- tame.” Bontekoe was in the island in birds in a state of nature without laws, kinson [4], undertook to demolish the 1618 and observed “eenige dod-eers-

160 American Scientist, Volume 100 © 2012 Sigma Xi, The Scientific Research Society. Reproduction with permission only. Contact [email protected]. en” unable to fly and very fat. Carré in a white dodo. Oudemans, however, Literature Cited 1668 writes of “celuy que les habitans thought that the male was deeper col- 1. Strickland, H. E., and Melville, A. G. The ont nomm? l’Oiseau Solitaire, parce ored and was depicted in certain paint- dodo and its kindred, or the history, affini- qu’effectivement il aime la solitude … ings that are now referred by Hachisu- ties and osteology of the dodo, solitaire and other extinct birds of the islands Mauritius, il ne ressem bleriot pas mal un Coq ka to young Mauritius dodos. During Rodriguez and Bourbon. London, 1848. 141 d’Inde, s’il n’avoit point les jambes the present century, at least three good pp. The copy examined in the Yale Uni- plus hautes. La beauté de son plum- representations have turned up that versity Library contains an inscription ‘To age fait plaisir à voir. C’est une couleur appear to Hachisuka to represent a sol- Professor Silliman with the kind regards of the authors” and a pencil note below changeante qui tire sur le jaune. La itaire-like bird, either white or brown, “Rec’d Octob 23—1848. The first part read chair en est exquise.…” with a very large head and conical and the second looked through within 8 In the next year Sieur D. B. (DuBois) beak. These he considers to be Reunion days”—Beneath this is the signature in ink wrote, “Solitaires: ces oiseaux sont solitaires, and, perhaps, overgeneraliz- of James W. Dana, doubtless added later. nommés ainsi, parce qu’ils vont tou- ing Leguat’s rather specialized concept 2. Hachisuka, Masauji. The dodo and kindred jours seuls. Ils sont gros comme une of feminine beauty, he depicts in his birds or the extinct birds of the Mascarene Islands. London, H. F. and G. Witherby, grosse Oye, et ont le plumage blanc, reconstruction the female as the more Ltd., 1953. 250 pp. The book is primarily a noir a l’extremité des ailes et de la beautiful white bird, largely from a wa- compilation, and at times somewhat naive; queue. A la queue il y a des plumes ter-color drawing discovered by Killer- its value to the non-specialist may, however, approchantes de celles d’Autruche, ils mann in Vienna. It is at first sight hard be judged from the fact that although one of the largest libraries in the New World ont le col long, et le bec fait comme to assimilate this drawing to a dodo, is available to the writer, he had to rely on celui de Bécasses, mais plus gros, les but if the white dodo lost weight and Hachisuka for the content of the works of jambes et pieds comme poulets d’Inde. the anterior sheath of its beak during Oudemans, Mortensen, Vivielle, and De- Cet oiseau se prend à la course, ne vo- the moulting season, such an identity hérain, and Killerman in preparing the lant que bien peu.” might be possible [6]. The decomposed above account. Strickland [5], who collated these wing of the Vienna drawing is very like 3. Voyage et avantures de François Leguat. Tome 1, 164 pp.; Tome 2, 180 pp. and index. accounts for the first time, dismissed that of the paintings of white dodos, David Mortier, Marchand Libraire, à Lon- Bontekoe’s use of the term “dod- and it seems inconceivable that two dres, MDC CVIT. The subsequently pub- eerse,” the ordinary Dutch for dodo, raphid birds, whitish, with the same lished English translation is inaccurate and as a confusion with the Mauritius wings, should have evolved in parallel misleading. bird, due perhaps to muddled mem- on the same island. Only skeletal evi- 4. Atkinson, G. The extraordinary voyage in French literature from 1700 to 1720. Lib. ory caused by Bontekoe’s ship being dence can now solve the problem, and Ancienne Honoré Champion, Paris, 1922. blown up and he alone surviving, unfortunately the volcanic caves on 147 pp. shortly after visiting Reunion. Strick- the island have not yielded bones. The 5. These extracts are all taken from Strickland. land concludes that the three other matter is of considerable general inter- The scientific name of the birds of Reunion accounts refer to a yellow or whitish est in view of the difficulties that might raises difficulties. De Selys Longchamp solitaire-like bird with a longer beak arise in explaining the origin of two named the solitaire of Reunion, on the basis of Carré and DuBois, Apterornis solitaria. than the Reunion bird. sympatric raphid birds on the same The generic name is preoccupied, but if a It is very curious that Carré’s and island. These difficulties are, moreover, single species, a white dodo, existed on the DuBois’ accounts, which have really presented in a more acute form by island it becomes Raphus solitarius (DeSelys confused the issue if only a single spe- the probable former presence of three Longchamp) or Ornithaptera solitaria if generic distinction is desired. If two birds are cies was present, both begin in a way species of giant tortoise on Rodriguez involved, the last is the name for the soli- reminiscent of the standard opening and of even more on Mauritius. A re- taire; Hachisuka has supplied Victoriornis of any chapter in a mediaeval bestiary. examination of the available material imperialis Hachisuka for the white dodo. One can easily imagine a fabulous spe- of these animals using modern statisti- 6. Miss Martha M. Dimock, in copying the cies being introduced with the words cal methods might be interesting. Par- pictures here reproduced, came to the con- Avis solitaria vocatur guia solitaria est.… enthetically, it may be mentioned that clusion that the two left-hand paler birds might well be of the same species. This sort Some years after Strickland wrote, Peter Mundy in the seventeenth cen- of testimony is of some interest in default of four very definite paintings of white tury wrote of the wingless birds of the direct observation, because in moving the dodos, by Pieter Withoos and by Pi- Mascarenes and of a wingless rail that hand in accordance with the old drawing eter Holsteyn, Sr., were identified in he, but no one else, observed on As- one probably gets nearer to the original bird than in any other way now possible. various European collections. It be- cension Island—”A question may bee 7. Peter Mundv’s travels. Hakluyt Soc. Ser. II, came evident that a dodo with whit- Demaunded how they should bee here 46, 335 and 78, 83. ish plumage, very decomposed, yel- and not elsewhere, beeing soe Farre lowish wings and plumose tail, ac- From other land and can neither Fly tually existed. It was clearly distinct nor Swymme; whether by mixture off from the Mauritius dodo in its color kindes producing straunge and mon- and less hooked bill, and presumably strous Formes, or the nature off the Cli- For relevant Web links, consult this could only have come from Reunion. mate, ayre and earth in alltring the First issue of American Scientist Online: Some authorities, notably Rothschild, Shapes in long time, or how” [7] When http://www.americanscientist.org/ have therefore suspected the presence another English traveler asked a like issues/id.95/past.aspx of two white forms on Reunion, one question of himself off the Galapagos dodo-like, the other solitaire-like. Oth- Islands nearly two centuries later the ers, notably Oudemans, suppose only answer proved to change the ideas of one species to have existed, and this the whole world. www.americanscientist.org © 2012 Sigma Xi, The Scientific Research Society. Reproduction 2012 March–April 161 with permission only. 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