Outlines of the Development of the Tuatara, Sphenodon (Hatteria) Punctatus
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Outlines of the Development of the Tuatara, Sphenodon (Hatteria) punctatus. By Arthur Demly, D.Sc, Professor of Biology in the Canterbury College, University of New Zealand. With Plates 1—10. TABLE OF CONTENTS. TAGE 1. PREEACE . .1 2. INTRODUCTORY REMARKS . .4 (a) On the Habits of the Tuatara . .4 (A) On the Time occupied in Development, with List of Embryos classified and arranged in Chronological Order . 9 (c) On the Structure of the Egg . .11 3. SYSTEMATIC ACCOUNT OF TIIE STAGES IN DEVELOPMENT . 14 4-. SUMMARY AND DISCUSSION OE PEINCIPAL EMURYOLOGICAL OBSERVA- TIONS . .60 («) The Formation of the Germinal Layers . .60 (6) The Formation of the Foetal Membranes . .05 (c) The Modelling of the Body, and the Foundation of the Princi- pal Systems . .69 (d) The Embryonic Colour Markings . ,77 5. LIST OE REFERENCES . .79 6. DESCRIPTION OP FIGURES . .SO 1. PREFACE. SHORTLY after my arrival in New Zealand my friend Pro- fessor G. B. Howes, LL.D., F.R.S., urged me to undertake the investigation of the development of the Tuatara (Sphe- nodon punctatus), but for various reasons I decided at that time not to do so. Some time afterwards, however,while examin- VOL. 42, FART 1. NEW SER. A 2 ARTHUR DENDY. ing the parietal eye in some embryo Australian skinks, I was struck -with the existence of an optic cup, apparently similar to that of the paired eyes. This seemed so startling that T resolved to attempt to obtain Tuatara embryos for the purpose primarily of studying the development of the parietal eye. Knowing that Stephens Island, in Cook Straits, on the recommendation of the Australasian Association for the Advancement of Science, bad been proclaimed a reserve for the Tuatara, it occurred to me by a happy inspiration to address a letter to the lighthouse keeper, then entirely unknown to me even by name, asking bis assistance in the matter. I wrote to him first on July 3rd, 1896, and most fortunately I found in Mr. P. Henaghan, the principal keeper on Stephens Island, an ardent enthusiast, who threw himself heart and soul into the work in the interests of science; and I cannot sufficiently express my gratitude to him for the magnificent supply of material which he obtained for me, and also for the valuable information which he gave me from time to time in his in- teresting letters regarding the habits of the Tuatara. Stephens Island being now a reserve, it was of course necessary to obtain permission from the New Zealand Govern- ment to collect, and this, through the kind assistance of Sir James Hector, F.R.S., was successfully accomplished. To the Hon. the Colonial Secretary and to Sir James Hector I also wish to express my gratitude for their courtesy. I had proposed visiting Stephens Island myself during the breeding season, which was supposed to begin about January, but pressure of other business made it very difficult for me to do so, and I ultimately resolved to trust entirely to Mr. Henaghan for the supply of eggs. There is regular com- munication between Stephens Island and the mainland only once in six weeks, so tlicit considerable difficulty and delay were experienced in sending the eggs, in consequence of which a considerable number perished. Several lots of the eggs were sent packed in moss or lichen in tin cans, but they were very liable to go bad in this packing if delayed OUTLINES OE THE DEVELOPMENT OF THK TUATAEA. 3 on the voyage, and subsequently we found that much the best way was to pack them, only a few together, in tin cans filled with the coarse brown sand which occurs on the island. Thus packed they travel admirably. I found it quite possible to keep the eggs developing, buried in damp sand after their arrival, but this was attended witli considerable risk, as they are very subject to the attacks of mould if kept too damp and without sufficient ventilation, while they readily shrivel up if allowed to get too dry. The first consignment of eggs was received about the end of January, 1897. Unfortunately the embryos all died, appa- rently from drying up of the eggs, before they reached Christ- tihurch. Nevertheless one advanced embryo, the only one as yet obtained of Stage Q, was removed from the egg in-suffi- ciently good condition to be of considerable value. I obtained no more eggs that summer, my arrangements being interfered with by a German collector, who visited Stephens Island about the new year, having obtained a recom- mendation to the keeper from the authorities. The only other eggs found that summer were forwarded to him, but I am informed that they perished in transit. The next eggs which I received arrived, greatly to my sur- prise, in July, 1897, a season of the year at which we did not at all expect them. They were only three in number, and con- tained very advanced embryos (Stage K.) in excellent condition. In November, 1897, the work began in earnest, and from that time to the present I have received consignments of eggs at more or less frequent intervals. Latterly I have requested Mr. Henaghan, when he finds a nest, to keep the eggs on the island, to continue their development, and send them on to me at intervals in small quantities, a plan which we find to work admirably, and which I regret did not occur to me before, as all the embryos thus obtained have been in much the same advanced stage in development, and some more of the earlier stages would have been more useful. The result of our operations, however, has been to secure a magnificent series of embryos, the majority of which hnvp heen 4 AETHUK DENDY. hardened in Kleinenberg's picric acid after removal from the egg, and preserved in alcohol. Those which were required for sections were stained with borax carmine, and cut in paraffin with Jung's sliding microtome. I found it desirable to use oil of cloves for clearing the embryos previous to embedding in paraffin, as this allowed of more advantageous examination of them as transparent objects than the turpentine method which I usually employ. It would, of course, be impossible in the time which I have had at my disposal to give anything like a complete account of the development of the Tuatara; and in this communication I propose only to give a general outline, with special reference to the formation of the germinal layers, the foetal membranes, the modelling of the body, the foundation of the principal r.ystems, and the classification of the embryos into stages; which will, I hope, be useful in future investigations either by others or by myself. This general account may be followed by more special memoirs dealing with the development of the particular organs. Already, indeed, I have sent home for publication one such memoir " On the Development of the Parietal Eye and adjacent organs." In concluding this preface I must express my sincere gratitude to Professor G. B. Howes for his kindly interest and encouragement, and for revising the proof sheets, and super- intending the execution of the plates in my absence from England. At my request he has undertaken to investigate the deve- lopment of the skeleton, and for this purpose I am forwarding to him a supply of material. 2. INTRODUCTORY REMARKS. (a) On the Habits of the Tuatara. Since I have not myself visited Stephens Island, it will be as well to give the following account of the habits of the Tuatara as nearly as possible in Mr. Henaghan's own words. OUTLINES OF THE DEVELOPMENT OP THE TTJATARA. 5 "Our island," writes Mr. Henaghan,1 "is densely covered with scrub of various kinds; the soil is good in most places, especially on the ridges; in the gullies the soil is of a light brown colour, largely composed of oxide of iron. The birds and lizards burrow into this soft soil, and one can often find both living peaceably in the same hole. There are three or four kinds of petrel frequenting the island, and if you were here now, which is their breeding season, you would be sur- prised at the numbers of them,—there is hardly a foot of soil but is undermined with their holes. Insects of various kinds are also well represented, and I think the lizards feed largely on them, especially beetles. I believe they also eat young birds,—in fact, I have seen them do it.s They, however, live a large part of the year without any food, keeping constantly to their holes. There are three or four kinds of lizards here alto- gether, the Tuatara being the largest; the others are very small. There have never been many lizards' eggs got here yet, though the Tuataras are very numerous." The breeding season apparently commences in November on Stephens Island, and each female lays about ten (10) eggs.3 On November 1st, 1897, Mr. Henaghan removed ten eggs from an individual which had probably been accidentally Crushed by a cow. These eggs, which were forwarded to me, were apparently quite ready to lay, but in the one which I Opened on their arrival I found no embryo, and the others unfortunately went bad subsequently, instead of continuing their development, as I hoped they might. The ground, Mr. Henaghan informs me, is so full of holes that one has to dig ever so many before finding a Tuatara, and then the probability is that it will turn out to be a male, these being far more numerous than the females.