< gf < CO a ae aE ae ae ee ic aie ra ee TOE ae Ce a a te rou Lan vy VWwvlyyi¥y Ae P = a . See ~. WWW e © Yee Se os eS age SSN SS RS SS Se eae ES SG =a 4 : — we : poe id Ww We y Wy GEE 235 PPR SS Sak BSS LOU LO GINA A PEN pe emenpen * e 3 ~~ — 4 te http:/www.archive.org/details/collectedscienti0Ogarruoft ail _T_E_——<— IN MEMORIAM. THE COLLECTED SCIENTIFIC PAPERS OF THE LATE ALFRED HENRY GARROD, M.A, F.R.S,, FELLOW OF ST. JOHN’S COLLEGE, CAMBRIDGE; FULLERIAN PROFESSOR OF PHYSIOLOGY AT THE ROYAL INSTITUTION; PROFESSOR OF COMPARATIVE ANATOMY IN KING’S COLLEGE, LONDON; PROSECTOR TO THE ZOOLOGICAL SOCIETY OF LONDON, EDITED, WITH A BIOGRAPHICAL MEMOIR OF THE AUTHOR, BY W. A. FORBES, B.A., FELLOW OF ST. JOHN'S COLLEGE, CAMBRIDGE; PROSECTOR TO THE ZOOLOGICAL SOCIETY OF LONDON. 0 HA LONDON: i R. H. PORTER: 6, TENTERDEN STREET, W. 1881. sf ) as F G0, vy ty, ’ ha oe PREFACE. In consequence of a very generally expressed wish amongst the personal friends of the late Professor A. H. Garrod, to possess some permanent memorial of him, it was decided, at a meeting of the Zoological Club held on May 4th, 1880, to appoint a Committee to consider the best means of carrying this idea into execution. As ultimately constituted, this Committee consisted of Profes- sors Flower, Schiifer, Bell ; Messrs. Sclater, Salvin, Balfour, Dobson, the late Mr. Alston, Dr. Giinther, and myself, I agreeing to act as secretary to it. After carefully considering the various forms that the proposed memorial might take, it was decided that the most appropriate and desirable one would be the publication, in a collected form, of all the papers published by Garrod in various scientific journals and periodicals, with a portrait and memoir of the author. The list of subscribers to the present volume, including, as it does, the names of nearly all the leading English biologists as well as those of numerous private friends, shows that this deci- sion of the Committee has been amply justified by the result. The work of arranging and editing the papers in the present volume has in the main fallen on myself, though Professor E. A. Schifer was kind enough to read through the proofs of the physio- logical part. As far as possible, the papers in each part have been kept in the order in which they were published: in a few cases, however, when several papers treating of the same or closely-allied subjects appeared at different periods, they have, where it seemed desirable, been placed together. The last paper in the volume, as explained in the biographical memoir, has not been published before anywhere. Misprints and other clerical errors of the original have been corrected in this reprint, and in a few cases, where it seemed advisable, foot-notes have been added by myself. The pagination of the original papers has been preserved by the use of marginal figures. le vi PREFACE. The octavo lithographic plates of the original papers have been _ redrawn by Mr. J. Smit for the purpose of this reprint, and they will, I think, be found faithful copies of the originals. The stones of the quarto plates being still preserved have, with the necessary alterations in lettering, been used again, as have also the plates of the three diagrams in the second part. The three plates illustrat- ing the first part have been redrawn, the first two being reproduced on a somewhat larger scale the better to suit the letterpress. The woodcuts in the text have been throughout printed from the original blocks, or from electrotypes of them, with the exception of that on p. 95, which was kindly drawn on wood, after the litho- graphed plate in the “ Journal of Anatomy,” for the purposes of this work by Mr. H. H. Johnston. The etching which forms the frontispiece speaks for itself, both as a work of art and as a portrait. The readiness with which Mr. Herkomer undertook to execute it was itself extremely grati- fying to the Committee, whilst the success of the result adds materially, they feel, both to the value and the interest of the pre- sent volume. The biographical memoir has been written by me, partly from my own personal acquaintance with the author and his work, partly from facts supplied me by his family. The index to Part IT, containing as it does the names of all the various species or groups of animals mentioned in it, will, it is hoped, prove useful as facilitating reference to the very numerous facts contained in these papers. Finally, on behalf of the Garrod Memorial Committee, I have to express our best thanks to the Committee of Publication of the Zoological Society, and to the Editors of the “ Journal of Anatomy and Physiology,” the “Ibis,” and “Nature,” for the very generous loans, for the purposes of this reprint, of the various stones or blocks of the plates and woodcuts illustrating the present volume which originally appeared in their publications, as well as to the Council of the Royal Society, who kindly allowed us the use of electrotypes of the diagrams published in the “ Proceedings” of that Society. W. A. FORBES. June 24th, 1881. LIST OF SUBSCRIBERS. P. S. Abraham, Esq. Prof. A. Leith Adams, F. R.S.; Dublin. E. R. Alston, Esq. Dr. J. Anderson, F.R.S., Calcutta. Harold B. Ansdell, Esq. Lieut.-Col. Godwin-Austen, F.R.S. F. M. Balfour, Esq., F.R. S., Cambridge. J. Barclay, Esq. Dr. L. Beale, F.R.S. Alfred Beddington, Esq. J. H. Beddington, Esq. Leopold Beddington, Esq. Prof. F. Jeffrey Bell. Dr. G. Bennett, Sydney. Graf von Berlepsch, Berlepsch, Hesse. W. T. Blanford, Esq., F.R.S. J. E. Blomfield, Esq. Rey. Prof. T. G. Bonney, F.R.S. Prof. T. W. Bridge, Birmingham. Dr. J. Lauder Brunton, F.R.S. A. G. Butler, Esq. W. H. Caldwell, Esq, T. D. G. Carmichael, Esq. Major-General H. Clerk. Col. E. H. Cooper. Dr. Elliott Coues, Washington. Prof. Curnow. Charles Darwin, Esq., F.R.S. Francis Darwin, Esq. The Duke of Devonshire, F.R.S. G. E. Dobson, Esq. Dr. Anton Dohrn, Naples. A. Dowsett, Esq. Prof. A. Milne-Edwards, Paris. D. G. Elliott, Esq., New York. J. Eric Erichsen, Esq., F.R.S. Sir J. Fayrer, F.R.S. Prof. W. H. Flower, F.R.S., Pres. Z.S. Dr. John Foo ~ J. 8. Forbes, Esq. Mrs. Forbes. Miss Forbes. W. A. Forbes, Esq. Dr. Michael Foster, F.R.S., Cambridge. W. Foster, Esq. William Francis, Esq. Rev. A. Fuller. Dr. Max Furbringer, Amsterdam. Dr. Hans Gadow. J. C. Galton, Esq.’ Dr. T. G. Garson. J. P. Gassiott, Esq. F. D. Godman, Esq. J. 8. Gordon, Esq. Col. J. A. Grant, F.R.S. Dr. A. Giintlier, F.R.S. Prof. A. C. Haddon, ve oe Dr. E. Hamilton. C. E. Haskins, Esq., Cambridge. Rey. E. Hill, Cambridge. G. Holding, Esq. R. E. Holding, Esq. The John Hopkins University, Baltimore. G. B. Howes, Esq. R. Hudson, Esq., F.R.S. Prof. Huxley, F.R.S. The Royal Institution of Great Britain. Dr. F. A. Jentink, Leyden. H. H. Johnston, Esq. Prof. E. Ray Lankester, F.R.S. The National Library of Ireland. The Manchester Reference Library. The Lord Lilford. J.J. Lister, Esq., Cambridge. R. Lydekker, Esq. P. T. Main, Esq., Cambridge. H. S. Marks, Esq., R.A. Dr. A. B. Meyer, Dresden, Prof. St. G. Mivart, F.R.S. H. N. Moseley, Esq., F.R.S. Prof. A. Newton, F.R.S., Cambridge. F. Nicholson, Esq. The Earl of Northbrook. H. F. Osborn, Esq., Princeton, U.S.A. W. Ottley, Esq. Prof. Owen, F.R.S. The Owens College, Manchester. T. E. Page, Esq. Vili LIST OF SUBSCRIBERS. Thomas Parkin, Esq. Herr A. von Pelzeln, Vienna. F. G. Penrose, Esq. Dr. W. Peters, Berlin. Henry Pollock, Esq. Prof. J. Prestwich, F.R.8., Oxford. Dr. Quain, F.R.S. E. P. Ramsay, Esq., Sydney. Capt. R. G. Wardlaw-Ramsay. Dr. B. W. Richardson, F.R.S. Briton Riviére, Esq., R.A. Prof. G. Rolleston, F.R.S8., Oxford. Rey. Eric J. 8. Rudd. Mrs. G. de la Rue. Prof. Rutherford, F.R.S. O. Salvin, Esq., F.R.S. J. E. Sandys, Esq., Cambridge. Howard Saunders, Esq. Prof. E. A. Schifer, F.R.S. P. L. Sclater, Esq., F.R.S. J. Scully, Esq. A. Sedgwick, Esq., Cambridge. H. Seebohm, Esq. R. B. Sharpe, Esq. J. Smit, Esq. B. Woodd-Smith, Esq. Prof. Stanton. H. Stevenson, Esq. Gibson Stott, Esq. Rey. C. Taylor, Cambridge. W. B. Tegetmeier, Esq. Prof. G. Thane. Oldfield Thomas, Esq. Trinity College, Dublin (per Dr. E. P. Wright). , Rev. Coutts Trotter, Cambridge. Prof. W. Turner, F.R.S., Edinburgh. Hugh Turner, Esq. W. Turner, Esq. The Lord Walsingham. Mrs. Eliza Ward. Prof. M. Watson, Manchester. W. F. BR. Weldon, Esq. G. F. Westerman, Esq., Amsterdam. H. T. Wharton, Esq. Rey. W. 8. Wood. J. Woodgate, Esq. Dr. E. P. Wright, Dublin. Prof. Yeo. . A, H. Young, Esq. The Zoological Society of London, BIOGRAPHICAL NOTICE. Any account of the life of the author of the papers contained in the present volume must be brief. Not only was his career cut off at a time when most men are but just entering on life, but it was also, in some senses of the word, uneventful. Born and educated under circumstances which obviate any of those struggles which have made the lives of many naturalists interesting, his subsequent life was one series of unbroken successes. Nor was it marked by any of those episodes of foreign travel which have fallen to the lot of many other scientific men, before they have settled down to a life of work at home. Alfred Henry Garrod was the eldest child of- Dr. A. B. Garrod, F.R.S. He was born in London, on May 18th, 1846, at No. 9, Charter- house Square, where his father had commenced to practise as a phy- sician a few years before. From 1856 to 1860, his father having mean- while removed to Harley Street, he was being educated in general subjects at All Souls’ Grammar School, Regent’s Park, and after the latter date entered University College School, Gower Street. At both these schools the classical authors of Greece and Rome were at that time the chief subjects of study. For them Garrod never developed any special taste, though in mathematics and drawing he took a greater interest, gaining a prize for perspective drawing at University College School. Garrod’s life, however, in so far as it can be of any interest to those who were not members of his family and private circle, com- menced with his entry at University College about October, 1862. Here he first began his acquaintance with the Natural Sciences, by attending the lectures, amongst others, of Prof. Sharpey, on physiology, and of Prof. Oliver, on botany, as well as those of Prof. De Morgan, on mathematics. From the latter, no doubt, and from his earlier education in the same subject, was derived that predilection for mathematical and mechanical studies which x BIOGRAPHICAL NOTICE, evinced itself so strongly in his after life. Those who knew him at that time date the commencement of his scientific enthusiasm, which was always afterwards very marked, from the first term of his attendance at Prof. Sharpey’s lectures, and he himself always spoke of that teacher with affectionate and admiring regard. i In 1864 Garrod matriculated at the University of London, passing in the first class. In the autumn of the same year he began his career at King’s College, London, having gained one of the Warneford entrance scholarships there, notwithstanding the fact that a considerable knowledge of classics was required for that competition. In the summer of 1865, whilst still a student at King’s, Garrod succeeded in obtaining the first medal in Prof. Oliver’s course of botany, at University College, a success of which he was always afterwards very proud; for it probably proved to him that, conscious as he must have been for some time of original power and grasp, he also possessed that capacity for steady applica- tion without which the former gifts are so often useless. Still working with indefatigable energy, he succeeded in obtaining the first, second, and third year’s scholarships for medical students at King’s, the highest success he could attain there. He remained working at King’s College Hospital till 1868, in which year he obtained his Licentiateship of the Apothecaries’ Society. It was during this period that his interest in the subject of the circulation of the blood, to which he subsequently devoted so much time and work, commenced. In the summer of 1868, in company with a younger brother, he made a trip to Marseilles by steamer, touching _en route at. Vigo, Lisbon, and Gibraltar, and returning by the same way, after a ctuise of about eight weeks. This trip Garrod enjoyed greatly, especially his visit to Vigo and Lisbon, about both of which places he was enthusiastic afterwards. While on board ship he devoted a good deal of his leisure to working at questions connected with the temperature of the body. Three years afterwards he again visited Spain, going to Cadiz, Seville, and Gibraltar, in com- pany with a college friend, but these two Spanish trips, and a flying visit to Switzerland in 1869, were almost all his experiences of foreign travel, for which indeed he often expressed a positive dislike in after years, - In 1868 Garrod went to Cambridge. Though he had put his name on the books of Caius College, and indeed appears in the University calendar for that year as a member of the College, he BIOGRAPHICAL NOTICE. xi never actually entered there, having had awarded him in the same year an exhibition for Natural Science at St. John’s, the first offered in that subject by the College. He commenced residence at Cam- bridge in the October term of 1868. At that time the opportunities and methods of biological teaching at Cambridge were not so perfect as they now are, and with his already existing anatomical and physiological knowledge, Garrod soon found out that he could very well dispense with a good amount of the ordinary routine of College and University education. Indeed the greater part of his Tripos work was done by him at home, in London, during the vacation, for at Cambridge _he devoted himself largely to the enjoyment of the social life of the place. Hisscientific work there consisted more of original research © than of the usual course of study, for it was during his residence at Cambridge that he carried on, in great part, the series of experi- ments on the causes of the varying temperatures of the human body, the results of which were subsequently made public. The circulation of the blood, too, he studied energetically, by means of the sphygmograph and other appliances, which he improved in various ways by his great mechanical genius. Photography like- wise took up some part of his attention, and when the new chapel at St. John’s was opened, in 1869, he succeeded in taking, from the rooms of a friend overlooking the scene, an instantaneous view of the procession as it passed though the first court. It was whilst still an undergraduate that his first physiological papers were published in the “Journal of Anatomy and Physiology,” and in the “ Proceedings of the Royal Society.” He devoted consider- able time also to the study of zoology, working in the University Museum, where he laid the foundation of his knowledge in the subject which was afterwards to chiefly occupy his life. His maiden zoological paper “On the Telson of the Macrurous Crustacea” (infra, p. 93) was indeed written and published during _ his undergraduate career at Cambridge. His College had meanwhile not been blind to the ability of their undergraduate member, for in 1870 he was elected a founda- tion scholar, and in the “ May” examinations in Natural Science of that and the subsequent year, his name appears in the first class on each occasion. In December, 1871, Barco “went out” senior in the Natural Science tripos, his companions in the first class being three in number, R. Lydekker—who has since distinguished himself as a xii f BIOGRAPHICAL NOTICE, paleontologist in the work of the Geological Survey of India— Lewis, and Warrington, in the order given. At this time the office of Prosector to the Zoological Society of London was vacant, owing to the resignation of its former occupant, Dr. Murie. Through the influence of his Cambridge friends, Garrod was brought forward as a candidate for this post, to which he was elected, on June 20th, 1871, though he had not at that time finished his “ Tripos.” On the conclusion of that examina- tion he returned to London, and immediately began to devote him- self, with his accustomed energy and ability, to the adequate per- formance of the duties of his new post. He commenced his attendance at the Gardens a few days before the close of the year 1871. Up to this time, as already stated, Garrod’s work and interest in biological science had been chiefly physiological, and his knowledge of zoology generally little more than that of an ordinary “ Tripos” student. But thanks to the opportunities of his position, and his own genius and diligence, he was enabled in a very short time to sufficiently master the main outlines of the comparative anatomy of the highest Vertebrata—Birds and Mam- mals—to present his first paper to the Zoological Society, one written in conjunction with Mr. Frank Darwin, whom he had known at college, in little more than two months after commenc- ing his prosectorial duties. This paper was succeeded in the course of the year by four others. From this point indeed, zoology became the main study of Garrod’s life, though he still retained his interest in physiology. Indeed he had hoped, and intended no doubt, when he became Prosector, to carry on his physiological researches on a still larger scale, as evidenced by some experiments made during the earlier part of his work at the Gardens. But the accumulation of material, and the fascination of new lines of research, gradually but surely drew him away from further active physiological work, and after his appointment as Prosector, we find but three or four original papers from his pen on purely physiological subjects, the last being one read before the Royal Society in April, 1874. Physiology, indeed, was his “ first love,” and towards the close of his life, he often, in conversation with his friends or relatives, insisted that he was, primarily, a physiologist, and only became a zoologist by the accident of his being appointed to the Prosector- ship. Nothing gave him greater pleasure during his last illness than the fact that conclusions the same as some of his own most BIOGRAPHICAL NOTICE. xiii cherished ideas and discoveries, connected with the circulation of the blood, had been arrived at independently by an American physiologist (Keyt) with no knowledge of his previous work on the same subject. In his new departure, the anatomy of birds soon became Garrod’s favourite study. This was a subject that had not at all kept pace with the rapid advance made of late years in most other branches of biolegical science. With the exception of some im- portant papers by our countrymen, Professors Parker and Huxley, by M. Alphonse Milne-Edwards, and by the great German natur- alist, Johannes Miiller, little had been done either at home or abroad in this department of ornithology, since the decease of the illustrious German, Nitzsch. With the large amount of material at his disposal, Garrod was soon able to work out,on a far more extensive scale, many of Nitzsch’s observations, as well as to add a great number of entirely new facts. The myology of birds in particular attracted his attention, and about two years after his appointment to the Prosectorship, he drew up the paper on the classification of birds, in which the taxonomic value of the now-celebrated “ambiens” muscle was brought forward for the first time. This paper was read before the Zoological Society in the session of 1873-4, and pub- lished in their “ Proceedings”: an abstract of it, doubtless from his own pen, may be found in “ Nature,” Feb. 12, 1874, pp. 290—2. In November, 1873, Garrod was elected to a Fellowship at his College, the first time that such an honour had been given there _ toa Natural Science man; and in the summer of the succeeding year (1874), he was elected Professor of Comparative Anatomy at King’s College, London, in succession to Prof. Rymer Jones, F.R.S., and this post he continued to hold till within a few weeks of his death. A report of his introductory lecture to the evening class of Zoology at that institution in the winter session of 1874 may be found in “ Nature” for Oct. 8th of that year. Garrod had for some time past acted as one of the sub-editors to the last-named journal, and he continued to do so for some years. Many of the articles and reviews dealing with biological subjects published in the columns of that paper, during that period, were from his pen. In 1875 Garrod was appointed Fullerian Professor of Physiology at the Royal Institution. His connection with that establishment had commenced at least as far back as 1874, when he delivered, on Feb. 6th, an address on “The Heart and the Sphygmograph,” an - Xiv BIOGRAPHICAL NOTICE. account of which appeared in the number of “ Nature” for the 26th of that month. Again,in March of the following year (1875) he gave a short course of lectures at the same institution on “ Animal Locomotion,” in one of which he exhibited and explained an ingenious model of a boat propelled by a screw, constructed on a plan similar to that of the dorsal fin of the pipe-fish.* Another dealt with the action of the Horse, giving an account of Marey’s experiments with the graphic method on that subject, which was then exciting a good deal of attention in England, on account of Miss Thompson’s celebrated picture of the “ Roll-Call,” exhibiting at that time. Garrod commenced his duties as Fullerian Professor in the spring of 1876, when he delivered a course of twelve lectures on the “Classification of Vertebrate Animals.” In these and his subsequent lectures at the same institution, his great mechanical ingenuity and extraordinary fertility of resource in devising and carrying out experiments, stood him in good stead. For-he was enabled, by his numerous models and other simple though ingeni- ous contrivances, to illustrate or explain many of the phenomena of animal life, in a way that always instructed at the same time that it entertained his audiences. These powers, aided by very considerable fluency as an extempore speaker, and great facility of lucidly explaining even complicated topics to a general audience, soon gained him a reputation as an accomplished lecturer. Many of those who attended these or his other lectures, must remember some of his ingenious models, which were all the more admirable often, because of their great simplicity. Such were his arrange- ments to show the mechanism of the protrusion and the retraction of the claws in the great Carnivora, of the power of rolling-up into a ball possessed by the Hedgehog, &c. Other instances of his mechanical powers that might be cited were his models of the bird’s wing, of the mechanism of the gizzard in birds, and of the action of the lips of the Manatee when feeding, exhibited in illus- tration of his papers on those subjects before the Zoological Society. In the summer of 1875 Garrod delivered several of the Davis Lectures at the Zoological Gardens, choosing as his subjects the various greups of Ruminating Animals, the Camels, Deer, Ante- lopes, &c. This group of Mammals had for some time past been * Vide “Nature,” April 1, 1875, p. 429. BIOGRAPHICAL NOTICE. XV the subject of his close attention, for but little regarding their structure had been put on record by previous anatomists, whilst he, thanks to his prosectorial advantages and the living collection around him, was enabled to study them, with every facility, both in a dead and a living state. The most important of his scientific results, regarding these animals, were embodied in a paper “ On the visceral anatomy and osteology of the Ruminants,” read before the Zoological Society on Jan. 2,1877. In this paper, besides many important conclusions arrived at with regard to the classification of this group of animals, Garrod broached his views on a subject that for some time had been more and more impressing itself upon him. This was the inadequacy of the ordinarily used system of binomial nomenclature to indicate properly the new views on the classification of animals rendered necessary by the general adop- tion of the theory of evolution—an inadequacy that was the more evident to him from his previous knowledge of chemistry. It was his desire to devise, if possible, some system of formule which would enable the biologist to express, in one term, both the nature and the affinities of the creature it represented. This effort to give taxonomic conclusions a more exact expression had been already partly adopted in the formule used by Garrod in his various papers on the classification of birds, as well as in the lecture on “ Evolution and Zoological Formulation,” delivered at King’s College in 1874, and already alluded to. In that lecture he had selected a parti- cular group of birds, the Parrots, to which to apply. his views: in his paper on the Ruminants these are still further expounded, and worked out for that group of animals. Meanwhile Garrod had been steadily going on amassing facts regards the anatomy of Mammals and Birds at the Zoological Gardens, and hardly a scientific meeting of the Zoological Society passed without some paper from him, dealing either with some of his newly-discovered lines of research or with an account of the anatomy of some hitherto unknown form. Some idea of the amount of material that passed through his hands may be gained from the fact that at various periods in his career he dissected no less than five specimens of Rhinoceroses, belonging to three different species—an anatomical experience probably quite unique. He worked remarkably quickly, both with his scalpel and with his pencil—for he was no mean artist—but yet with such certainty and accuracy that a re-examination of his objects rarely rendered necessary any change in his original description. His anatomical XVi BIOGRAPHICAL NOTICE. work was as a rule directed rather to the discovery of new points bearing on questions of classification than to detailed description of forms only interesting for their rarity, and agreeing in most points with the typical species. His method was to examine as large a number of species of any particular group as possible, to note the characters in which they differed, and from these con- siderations to endeavour to arrive at some conclusions regarding their general position or mutual affinities. When the Royal Commission appointed by the Government examine into the question of vivisection was conducting its inquiries in 1875, Garrod was, with most of the other leading physiologists of England, examined before it, and a report of his evidence will be found on pp. 106—108 of the Blue-book that contained the results of their inquiries. Amongst other things, he narrated how he had, on one occasion, performed “ probably the largest operation in anesthetics” ever effected, by chloroforming a Giraffe. Garrod had been very anxious to utilise the opportunity afforded by having to kill one of these animals some time pre- viously, by taking tracings of the blood-pressure in its carotid arteries, which, after a considerable struggle with the great animal, much expenditure of chloroform, and a good deal of difficulty in properly fixing the necessary instruments, he succeeded in doing. In 1876, being then just 30 years of age, he was elected a Fellow of the Royal Society. At this time the expediency of preparing a general work on the “Anatomy of Birds” was suggested to Garrod by some of his friends. This work was to be, in his own words, “ exhaustive,” and to contain the results of all his previous papers and subsequent experience. For the completion of this he soon set to work with great vigour, and on two occasions he received sums of money from the Government Grant to aid him in the publication and preparation of this book, which, unfortunately, he was not destined to finish. In 1876 Garrod was appointed examiner in Zoology for the Natural Science Tripos at Cambridge, a post which he continued to fill for the next two years, and which again brought him into intimate connection with Cambridge life. The subject of his Fullerian lectures in 1877 was “The Human Form: its structure in relation to its contour.” In them it was his object to describe to a general audience those parts of the BIOGRAPHICAL NOTICE. xvi structure of the body which affect its external form in various attitudes. Several ingenious working models were devised espe- cially to illustrate these lectures, as well as a colossal papier-maché model of a disarticulated human skeleton. The attendance, more than half of which consisted of ladies, on this course of lectures was very large, and fully proved the success of his experiment. In the following year he chose as the subject of his last course “The Protoplasmic Theory of Life, and its bearing on Physiology,” in it explaining the nature of the modern cell-theory, and the conclusions deducible from it. These lectures were likewise numerously attended. . Meanwhile Garrod maintained as actively as ever his re- searches into the anatomy of Mammalia and Birds, and his pro- jected work on the latter group occupied much of his attention. The anatomy and classification of the enormous group of Passerine Birds especially attracted him, and he published four important papers on that subject in the Zoologica] Society’s Proceedings, pro- pounding in them a new division of the entire group, based on his own and others’ observations. About this time he took in hand the production of an English edition of Johannes Miiller’s cele- brated paper on the vocal organs of Passerine Birds. This had originally appeared in the “Abhandlungen” of the Berlin Academy; but, although containing results of the greatest im- portance, was hardly at all known in England. It was translated into English by F. Jeffrey Bell; and edited, with an appendix con- taining much additional information on the same subject, arrived at from his own researches, by Garrod. Early in 1879 it was completed and published, the publication having been undertaken by the delegates of the Clarendon Press, Oxford. Another piece of work that may be mentioned was the section on the “Ruminantia” in Cassell’s “ Natural History,” in which he still further elaborated his investigations into the natural history and anatomy of his favourite group. Being based on original observations and study of specimens, its value must not be measured by that of certain other portions of the same book, for it undoubtedly gives the best general account of this group of animals yet published in our language. During the three years’ cruise of H.M.S. “Challenger,” the naturalist staff had made a considerable collection of birds in spirit, which was especially rich in oceanic birds, particularly in the group of Petrels and Albatrosses (Tubinares). With the examination of b Xvili BIUGRAPHICAL NOTICE. a these Prof. Garrod was entrusted on the return of the expedition, and the specimens so obtained were the more valuable to him, ‘because they helped to supply him with what had hitherto been a great desideratum. Not content with his own labours, Garrod was always anxious to induce others to work at his favourite studies. His rooms at the Zoological Gardens became gradually a centre for many other young men, who either aided him in his own work, or carried out further researches under his directions, or at his instigation. Those whom he thus helped were not all zoologists, for many artists could also testify to their indebtedness to him for the help in material, or the instructions in questions of anatomy, which he always freely gave when in his power. The society of artists, indeed, had par- ticular attractions for Garrod, and until his death he was a member of the Arts Club in Tenterden Street. These multiple and various occupations, added to an almost feverish activity of temperament, and an entire forgetfulness of self, proved, alas! before long too much for Garrod’s physical strength. Perpetually occupied in his scientific work, with new ideas opening out as every day brought fresh material and know- ledge, taking little or no holiday, and with the Italian Opera during the season as almost his sole relaxation, his health at last gave way. With hardly a premonitory symptom, Garrod was seized, in the early part of June, 1878, with a severe attack of pulmonary hemorrhage, which completely prostrated him for some time, and to those who knew caused apprehension of the gravest kind for the future. However, after some time he rallied somewhat, and became again capable of carrying on his prosectorial work. Although urged to leave England and try the remedial effects of a better climate, his love of London and work induced him to remain, heedless of the representations of his friends, and through the rest of the summer and autumn he continued to work in his usual enthu- siastic manner, though his health was obviously failing him. In the October of that year he delivered, without a note, the intro- ductory address at the commencement of the winter session of the King’s College Medical School, and in the course of the next month he paid a visit to Cambridge, to take part in a meeting of the Fellows of his College to decide on the new Professorships, which it was proposed that institution should endow. Garrod himself was strongly in favour of founding a Professorship of General Biology—a chair, as he insisted justly, greatly wanted in BIOGRAPHICAL NOTICE. xix the University, but his opinion was not shared by the majority of his colleagues, and, as he had not prepared any very definite scheme as to the functions of and qualifications for the new Professorship, his proposition fell through. In December, 1878, Garrod paid his last visit to Cambridge, when he came down to examine for the final part of the Natural Science Tripos. His friends there were all alarmed at the marked change in his looks and health now manifest. Yielding at last to the pressure put upon him by his family and friends, Garrod was induced to leave London and try the more genial climate of the Riviera for the winter. Accordingly he left England about the middle: of December, and went, accom- panied by one of his brothers, to Mentone, where he stayed till the end of January. During this period he was for some time confined to his bed, and the change, partly because the season was wet and dull there, did but little good. Returning to London, he still continued, whenever strong - enough, to work, visiting the Gardens when the weather was fine enough to allow him to do so, and directing from his home, as far as possible, his work there. But his health was gradually failing, and he grew almost daily weaker and weaker, as the phthisis and its accompanying maladies increased. Unfortunately for himself, he was far too well aware of his own danger, and indeed from the first seizure, or shortly afterwards, he considered his own case as practically hopeless. Nevertheless, he never ceased for an instant from his zoological studies. The subject of the conformation of the trachea in the different groups of birds was in particular at this time attracting his attention. At these windpipes, being con- veniently-sized objects, he constantly worked, whenever able to be up, at his own home, and one part of his observations, dealing with the trachea of the Gallinz (infra, p. 477), he sufficiently completed to be enabled to publish it. This was his last published contribution to that anatomy of birds which he had advanced so greatly and loved so well. : In the summer of 1879 there was for some time an apparent change for the better in Garrod’s condition. He seemed stronger and more sanguine than he had been for some time past, and was enabled to go again to his rooms at the Gardens with some regu- larity. But it was only temporary. The last time, so far as the writer is aware, when Garrod visited the Gardens, was about the middle of August. Shortly afterwards, about the commencement of September, he became again worse, and rapidly began to sink. xx BIOGRAPHICAL NOTICE. Almost to the last he continued to work, and only a few weeks before his death busied himself in correcting the proofs of his paper “ On the Brain and other parts of the Hippopotamus,” which had been read at the concluding meeting of the Zoological Society in the preceding June, but this, his last scientific paper, he was not himself destined to see published. During the later phases of his illness, which he had throughout borne with exemplary patience and courage, laryngitis, causing a nearly total loss of voice and great difficulty in swallowing, had developed itself, together with a severe cough. Under this complication of disease, he at last succumbed, in the perfect possession of all his faculties, though physically fearfully weakened, surrounded by his family, and re- eretted by all who had ever known him, on October 17th, 1879, aged 33. Of the worth of Garrod’s scientific work, the papers contained in the present volume must be the criterion, and the full value of some of them, as he himself was the foremost to believe, will only in all probability be properly appreciated in the future. This is not the place or the occasion to attempt to form a final conclusion on that score. His own confidence in his physiological work re- mained unshaken, and the partial confirmation of some of his ideas that he lived to see was, as we have already said, a cause of the most lively pleasure to him on his death-bed. Whatever may be the verdict of posterity on Garrod’s physio- logical work, no doubt can exist as to the value of his zoological labours. The facts alone recorded in his various papers on this subject must always remain as a great and incontrovertible addition to our knowledge of the highest Vertebrata. His published papers only represent a portion of the work he had done in zoology, for he had accumulated in notes and drawings; as well as in his memory, an immense amount of information on the structure of both Mammals and Birds, parts only of which had been utilised in his various papers. These notes and drawings fill several volumes of note-books, and have fortunately been preserved intact. Of his work on the “ Anatomy of Birds,” which was originally to have been published in three fasciculi, the MS. of the two portions com- menced has heen also fortunately preserved. As originally planned, the first fasciculus of this book was to contain a detailed account of the anatomy of the common fowl, as a type of birds in general. The second fasciculus was to be devoted to a comparative account: of the “ soft parts” of the different families of birds in systematic BIOGRAPHICAL NOTICE. Xxi order, whilst the third was, we believe, to have been devoted to the osteology, and a general réswmé of the results arrived at as regards classification. Of these fasciculi, the MSS. of the first and second were well advanced at the time of Garrod’s death ; the first part indeed is nearly completed, whilst the second is about half done, and it is hoped that at no very distant date these may be completed and published by his successor in the Prosectorship. Besides these MSS. one or two incomplete papers have been left, though hardly any sufficiently finished for publication, except the one herein contained, and now published for the first time, on the anatomy of the Diving Petrel (Pelecanoides). There are in addition a very large number of detached notes and drawings, many of the latter, depicting the conformation of the trachea in different groups of birds, being the more interesting _ as having been made during his lingering illness when not too weak for such occupation. Of his zoological papers indeed, the orni- thological ones must probably, on account of their more novel character, and as affording entirely new data for the solution of the various problems connected with the classification of Birds, which he revolutionised, be considered of the greater importance. No future worker in that group can neglect the facts or ideas con- cerning it that we owe to Garrod, and they alone suffice to put his name in the very first rank of those who have ever studied these creatures, and to stamp his work on Birds as truly “ Epoch- machende.”* A final word may be said on Garrod’s character as a man. The universal regret which his premature death has caused amongst all who knew him, whether intimately or casually, naturalists or not, is the surest sign of the esteem in which he was held. Unselfish and generous, he was always ready to help anyone as far as possible, with advice or material. Always cheerful, and with a wide range of information and interest, he was a companion of whose society one never wearied. A man of strong character, with great energy and pronounced views on many subjects, he yet did not make enemies. Those who knew him best loved him most, and feel deeply how little is the chance of their meeting with his like again. * A more detailed review of Garrod’s ornithological work, by the editor of the present volume, will be found in the “Ibis” for 1881, pp. 1-32. ne é eee 7 if cs OU a CONTENTS. PART I. On the cause of the diastolé of the ventricles of the heart .....ssesceeees On some of the minor fluctuations in the temperature of the human body when at rest, and their cause)... 0... se cc ee cece ep cece sateen escces On the relative duration of the canned parts of the edial sphyemoemeh trace in health. (Plate L.).. = : ave awaken ia eigdiccorecti-tenesngs feorn.the human cheet-wall "(Plate IL) enn aeie o1018 On the construction and use of a simple cardio-sphygmograph .........+++- On the mutual relations of the apex cardiograph and radial spygmograph Beate salak ioe: GF tha tseapersture ofthe air to tint off the, body Sgideied wee On pulse-frequency, and the forces that vary it ........+-seeeceeeeeceeees fae We De, Hee eerie he otreeed AEN BOE = wersee te usvsiee On sphygmography ......-2+.+eeeeeeese dieivo/ sale abSje 914 Sxe Oe nainene ie ths desires of nerve Biers epi tas esha « Aaes sion On some points connected with the Srealation: of the blood, mek a MOS a study of the sphygmograph-trace. (Plate III.) .. Jae woagite aelderey PART Il. ies hye telecon uf the. Macrurous Crustaces. . Seb kalgudts OV eicw eben sii cosmos Notes on an Ostrich lately living in the (Zoological) Society’s Collection Javan On the mechanism of the gizzard in Birds . Jey Su eae oe Notes on the anatomy of the Huia Bird (Heteralocha ala ts is sus de Note on the tongue of the Psittacine genus Nestor .. pete Sine cba s Note on some of the cranial peculiarities of the Woodpeckers Pe Cet Note on the placenta of the Hippopotamus ....... Vo Ve welc WS eine Cnideloa On the order Dinocerata (Marsh) ......-..- Wes tila we be dace sa ee entee barat ie Of the value in classification of a peculiarity in the anterior margin of the nasal bones of certain Birds ....... ts seks ee On the visceral —— of the Sipabuas athinccaros (UCmraleehinds suma- trensis) . : E On the tress of ‘has Sumetvaa Tiassa (Garatiehdens: Sausatttnniah: (Plate LV.) << siste cin we cect ewes see Sue de ses ce disewes co sweeeeee vic Swbse On the death of a Rhinoceros in the Society’s Gardens, and on some points in its anatomy ....... ecenee veescces On some points in the viecetal wiiaboaly of the Welcatishiton of the Sucailenliade (RAMMDCATOS SORAAENS) Foc .on ces ahaa Ss Coed teh scleatentavenewec's On the Tenia of the Titacadeos of the il cate (Plagiotanta ‘Pinaion; GEGES) \« arcinha’arachacre niall eee are ame Ste Sed ake Beles win dia ere slenlale oeiae Se ss 146 151 r XXiv CONTENTS. _Notes on the anatomy of the Binturong (Arctictis binturong) cose. ecee eee Note on the anatomy of the Binturong (Arctictis binturong) .....0+.0.000% On the cause of death in a Black-faced Kangaroo oe melanop).. On ‘the carotid arterids of Birds’. ¢.-s0s des eee tuisbink ween ee eee ee On some points in the anatomy of Steatornis ..... o 810 916 ss 80 ae ele 60 On certain muscles of the ee of Birds, and on eek nile in Gindeitioatioe (Plate V.) .. er <0.04 epee R OSs Sa anew sipe eeioe eae nstes ce Note on the cecum at Canis cancrivorus . On the visceral anatomy of the Ground Rat (Anlaseies pokabaouiaiie On some points in the anatomy of the Columbe........ ae vesewe Notes on two Pigeons, lanthenas leucolema and Wag the caine paleherrina. On the “showing-off ” of the Australian Bustard (Zupodotis australis)... Further note on the mechanism of the “ showing-off” in Bustards .........- On some points in the anatomy of the Purrots which bear on the classification of the suborder. (Plates VI. dnd VIL) i...0ss.seccccccseusvecstevc Notes on the anatomy of certain Parrots .......0..seeeees: See Note on the absence or presence of a gall- -bladder in the Sanntly of the Parrots 2.44 iieevey wee estas diye. s Stee es saind ate daremite . On the Kangaroo called Halmaturus tchiosnd is D’Albertis, ead its aihaatibea (Plates VIII.—X.) . Ata On a point in the isacdailian of tlie’ Bird’s wing. ora 9469/9 a DIE Be Wiaiwhte garantie On the form of the lower larynx in certain species of Ducks............. Ares On the form of the trachea in certain species of Storks and Spoonbills ...... On the disposition of the deep plantar tendons in different Birds............ On the Hyoid bone of the Elephant .. sia ieeesuhe an Fene Report on the Indian Elephant which died $i in the andaas J uly 7th, 1875 .. Notes on the Manatee (Manatus americanus) recently coe in the Society’s Gardens. (Plates XI.—XIII.) .......... wale ede veiv eee On the cecum coli of the Capybara (Hodrockeires eapybara) . Lieticses eat On a peculiarity in the carotid arteries, and other ee in the anatomy, of the Ground-Hornbill (Bucorvus abyssinicus) .. On the anatomy of Chauna derbiana, and on the ° systematic ‘position € the Palamedeide. (Plates XIV.—XVIL.).. {xcteeseiteiipath aie elt alae Na On the anatomy of Aramus scolopaceus .... . rrr rey Notes on the anatomy of Plotus anhinga. (Plates XVII. ~XxX).. Note on points in the anatomy of Levaillant’s Darter (Plotus Jevaillawtt) <. Notes on the anatomy of the Colies (Colius) . scotia ean creates 60 avs iesae On some anatomical peculiarities which bear poet the “iat divisions of the Passerine Birds. Part I. (Plates XXI.—XXVL_) ....ccccecceesvees Notes on the anatomy of Passerine Birds. Part II. ........ Baked wicyew tise ds Notes on the anatomy of Passerine Birds. Part III. (Plate XXVII.)...... Notes on the anatomy of Passerine Birds. Part IV... ....220eoeeeseereces On the Chinese Deer called Lophotragus michianus ey Mr. Swinhoe. sai XXVIII). Notes on the risheoeh anatomy of the ‘Ramiianie with a sguamostion sognediog a method of expressing the relations of species by means of formule ...... Note on the solid-hoofed Pigs in the Society’s collection .:......++seeeeees On the mechanism of the intervertebral substance, and on some effects of the erect position of Man ....... 2 3.6 eaters Notes on the anatomy of the Musk- ‘isan (iloeshas moins) -. chamies Notes on the anatomy and pines: position of the genera Thitnaccbin cae AEEEGIS 8.65 Sava 6:0 + vais Note on an anatomical naculintity 3 in claus Storks aa a's nn eh DOS eRe 415 421 CONTENTS. Notes on the anatomy of the Chinese Water-deer (7: ee reset Note on the possible cause of death in a young Seal . On the systematic position of the Momotide ..... Note on the gizzard and other organs of Cirpephugh icinaae Notes on the anatomy of Tolypeutes tricinctus, with remarks on other (hie dillos . z Notes on the ‘viaceua niahaaly of Kasai ‘ithe Sad ‘of Nycterentes p adieee- On the trachea of Tantalus Teculatie ‘on of Venetia’ IN THE TEMPERATURE OF THE HUMAN BODY. No. V.—From 10.30 p.m. till 12 night. 98° 99° 100° Sitting in a room (temp. of air 58°F.) all the time. Warmly clad till 11, when stripped in two a minute into a colder room. At 11°45 put on several flannel things, which had been warmed by the fire, and sat in front of a warm fire. Took sphygmograph-trace from right superfi- cialis vole at 10.40 and at 11.10. Tried to do so at 11.40, but could not get any indication, from the smallness of its pulsation. At 12 the pulsa- tion was as great as at 10°40. 98° 99° Sitting in a room (temp. of air 59°F.) from 9.30 until 10.40, quiet, cool, and warmly clad. From 10.40 till 10.55 moving about in the same room. Stripped at 10°55, and nude in two minutes. Remained nude until 11.24, when got to bed, and remained there for the rest of the time. minutes, so nude at 11.2. At 11.20 went for half Page 422, Page 423. 10 ON SOME OF THE MINOR FLUCTUATIONS No. VII.—From 11.10 p.m. til 11.55 P.M. Standing in aroom (temp. of air 53° F.) from 11 until 11.25. Fully clad until 11.9, when stripped, and nude at 11.10. Continued nude until12. At 11.25 seated, and remained so until 12, on a bed. At 11.40 put feet in water from 110°— 114°, above ankles, and re- mained thus rest of time, main- taining the heat of the water. Chilly when feet in bath, not 100° before. At 11.523 contracted limb muscles tonically, and maintained them so until 11°55. © Indicates temperature of pectoral region, two inches above nipple, taken with spiral thermometer, for five minutes. No. VIII.—From 11.25 p.m. till 12.40 night. 99° Standing in a room (temp. of air 58° F.) from 11 until 12, and sitting during the rest of the time on a bed. Fully clad until 11.50. Nude from 11.52, and remained so. Feet a little cold at 12.20, and put them into hot water (108°<114°) at 12.21, gradually in- creasing the heat of the water. Kept feet in water, above ankles, until 12.40. On adding more hot water and putting feet in it chills followed. No. [X.—From 9.15 a.m, #11 10 a.m. 95° 100° Sitting all the while in a room (temp. of air 52° F.), not far from an ordinary fire. Felt cold all over during the time. Reading. At 9.30 turned to the fire and put feet on the fender, having been previously quite at the side of the fireplace. As feet got warm, hands, which were previously warm, became cold, Clad in winter clothes. IN THE TEMPERATURE OF THE HUMAN BODY. 11 No. X.—From 11.10 a.m. tll 12.40 p.m. 99° 100° Temperature of air 62° F. A cloudy, breezy day. At ET oR 11 walked about 200 yards on to a beach, and sat down on the shingle at 11.5, where there was a slight side breeze. Hands and feet a little cold. Sun covered by clouds until 11.35, after which it be- gan to shine; immediately after which began to feel warm, and continued to get warmer until 12.7, when at 12.7 a cloud covered sun until 12.11. During time sun covered, several chills came over body. Walking in sun from 12.16 onward. 99° 100° Clad in thin merino next skin and summer clothes. No. XIl.—F rom 3 p.m. till 6 P.M. Temperature of air 66° F., slowly diminishing to 64°F. Sitting on a beach from 3 until 5, after a dinner at 2.15-2.45. A slight face breeze. Intheshade. Warm until 4.15, when feet: began to get a little cold, and by 5 so cold that obliged to move about. At 5 began to walk slowly, and had to go up several steps. At 5.20 began to walk briskly. Began to perspire at 5.25. Continued walking, per- spiring until 6. Clad as in last. To explain these Tables :— The actual temperature of the body at any given moment must be the resultant of (1) the amount of heat generated in the body, and (2) the amount lost by conduction and radiation. (1.) The source of heat in the body is not considered in this paper ; and no more will be now said of it, except that there is every reason to believe that it is not in the skin itself, and that, for the short periods through which each observation was made, it is approximately uniform. (2.) The loss of heat from the body is modified by changes in the skin and by changes in the surrounding media; and these two are mutually dependent. It has long been known that cold contracts and heat dilates the small arteries of the skin, respectively raising and lowering the arte- rial tension, and thus modifying the amount of blood in the cutaneous. capillaries. But modifications in the supply of blood to the skin must alter Page 424, Page 425. 12 ON SOME OF THE MINOR FLUCTUATIONS the amount of heat diffused by the body to surrounding substances ; and so we should expect that by increasing the arterial tension, thus lessening the cutaneous circulation, the blood would become hotter from there being less facility for the diffusion of its heat, and that by lowering the tension, thus increasing the cutaneous circulation, the blood would become colder throughout the body, from increased facility for conduction and radiation. That such is the case is proved by Tables I, II, III, IV, V, and VI, where, by stripping the warm body of clothing, in a cold air, when the tension was low (as in Tables IV, V, shown by the sphygmograph- trace), the temperature and tension rose, at the same time that the surface became colder. In Tables I, If, IIIf, IV, V, and VI, by covering the nude body with badly conducting clothing, when the tension was high, the surface- heat soon accumulated sufficiently to cause a sudden reduction of arterial tension, commonly called a glow, and a rapid fall in the tem- peratures, from the larger amount of blood exposed at the surface of the body to the influence of colder media. Changes in the arterial tension are easily recognised by the subject of experiment, from the sensations they produce; a feeling of warmth followed by a shiver, or a shiver itself, generally shows that the tension is lowered, while the opposite effect follows a rise in the tension; and this can be generally confirmed by the sphygmograph-trace. A bounding weak pulse shows a low, and a small thready one a high tension. - We know, from the observations of Davy and others, that by reducing the tension in one part of the body the tension of other parts is lowered; thus by placing one hand in hot water, a thermometer in the other rises. In Tables VII and VIII, it is shown that by putting the feet in hot water (at 110° to 115°) the lowering of the tension was so great that the amount of heat lost into the air considerably exceeded that gained'to the body from the water, so that the temperature of the body began to fall directly, and decreased considerably ; and it was noticed that on adding more hot water chills were produced, which was the same as the effect of first putting the feet in the water. By covering a small part of the body with a bad conductor, the tension of the whole body soon falls, from the accumulation of heat in the covered parts causing a lowering in the tension generally, and a consequent greater carrying away of heat. In this way the fall after- sitting down on a bad conductor when nude can be explained (Table VII). A glow is felt in the skin directly upon short muscular movement, as stooping, and the temperature falls at the same time, asin Table IV, between 11.45 and 12.20, and in Table XI, between 5.0 and 5.15. In the latter case the muscular movement was carried to such an extent IN THE TEMPERATURE OF THE HUMAN BODY. 13 that the loss was made up for by the increase of heat from en muscu- lar movement. Simply heating the feet lowers the tension and temperature OR: as in Table IX and in Table X. The passage of a cloud before the sun seems to have acted by reducing the loss of heat, as the tempera- ture rose at the time. Further confirmation of the facts stated as to the modification of Page 426. arterial tension may be found in Marey’s work, De la Circulation du Sang, published in Paris in 1863. In that book the author ascribes the uniformity of the heat in the internal parts to the same cause as the author of the present paper ascribes the variations. The fact observed by Dr. W. Ogle in the St. George’s Hospital Reports for 1866, and by Drs. Ringer and Stewart in a paper read before the Royal Society this year, that the temperature falls at night, and is lowest at from 12 to 1 a.m., and begins to rise after that time, is simply explained on the theory given above ; for it depends on the custom of Englishmen going to bed at about that hour, and thus giving a large amount of heat to the cold bedclothes, which at first is expended in warming the sheets, &c., while later on in the night the bedclothes are warm, and therefore the body has only to make up for the heat diffused. Other natural phenomena can be similarly explained. Thus, on a cold day, the effect of sitting with one side of the body in the direct rays of a fire is to cause the other side to feel much colder than if there was no fire at all, because the fire lowers the tension over the whole body, and supplies heat to the full cutaneous vessels of one side, while the other side, being equally supplied with blood in the skin, does not % receive heat, but has to distribute it rapidly to the cold clothes, &c. P ee ae 8 ne oe ve Page 351. 14 ON THE SPHYGMOGRAPH TRACE. 3. ON THE RELATIVE DURATION OF THE COMPO- NENT PARTS OF THE RADIAL SPHYGMOGRAPH TRACE IN HEALTH.* (Piate I.) Tue graphic method of representing the various phenomena occurring in the body during life, which has been so much developed by MM. Marey and Chauveau of Paris, has placed within our reach great facilities for obtaining an accurate knowledge of the relations, in point of time, of mutually dependent physiological events, and the sphygmograph has become, among others, an instrument familiar to most interested in science. By means of this instrument a detailed and truthful record can be easily obtained of the modifications in the diameter of any superficial artery, and, as usually constructed, it is intended to be applied to the radial at the wrist. _ The traces to be referred to were taken with one of Marey’s instru- ments, as made by Breguet. The recording paper ran its whole length, 43 inches, in seven seconds, and thus, by counting the number of pulse-beats in each trace, and multiplying the number thus obtained by 8°57143, the rate of the pulse at the time the trace was taken “was easily found. The lever-pen was of thin steel, sharply pointed, and it recorded by scratching on highly-polished paper previously smoked. It is now generally agreed that in each pulsation of the radial sphygmograph trace, the main rise is the effect of the contracting ventricle sending blood into, and thus filling, the arterial system. This rise is followed by a continuous fall when the pulse is quick, but when slow, its continuity is interrupted by a slight undulation, convex upwards. The major fall is followed by a secondary rise, not so considerable as the main one, but more marked than any other, and this secondary rise is evidently due to the closure of the aortic valves preventing further flow of blood heartwards. The two points therefore, the commencement of the primary and of the secondary rise, may be considered to mark the beginning of the systolé of the heart, and the closure of the aortic valve respectively, as far as they influence the artery at the wrist; and the interval between these two events may be called the first part of the arterial sphygmo- * “Proceedings of the Royal Society,’ XVIII. pp. 351-4, Pl. II. Read May 19, 1870. ON THE SPHYGMOGRAPH TRACE, 15 graph trace, while the interval between the beginning of the secondary Page 352. rise and that of the succeeding primary one constitutes the second part of the same trace. In 1865, Prof. Donders* published the results of experiments to determine the relative duration of the first and second part of the cardiac revolution with different rapidities of movements of the heart; taking as his data the commencement of the first and second sounds respectively, and he came to the conclusion that, though the second part varied with the rapidity, the first part was almost constant in all cases. On commencing work with the sphygmograph, the author came to the same conclusion with regard to the trace at the wrist, but, on improving his methods of observation, he has arrived at a different result. The best means of insuring an accurate measurement of any sphygmograph trace is to project all the points desired to be com- pared on to one straight line, and this is done by fixing the trace on to a piece of board, which has another pointed lever attached to it, with relations similar to those of the lever and recording apparatus in the original instrument. By this means lines can be scratched on the trace similar to those which would be produced by the instrument itself if the watch-work were not moving, and a result, as shown in Plate I. fig. 1. can be easily produced. The reason why this means has to be employed is, because the lever in the sphygmograph moves in part of a circle, not directly up . and down. > The ratio between the length of the first part of each pulse-beat in a trace and that of the whole beat was measured with a small pair ; of compasses, and from these the average was obtained, which thus ? eliminated, in a great degree, the variations produced by the respira- é tory movements, and also some of the clock-work imperfections. al For example, in fig. 1, the ratios in the several beats are :— 4 1:18 * | : 1-725 : 1°725 :1°775 : 1°725 es : 1725 : 1775 :18 * “ On the Rhythm of the Sounds of the Heart.” By F.C. Donders. Trans- lated in the “ Dublin Quarterly Journal of Medical Science,” Feb. 1868, from the “Nederlandisch Archief voor Genees- en Natuurkunde,” Utrecht, 1865. Page 353. 16 ON THE SPHYGMOGRAPH TRACE. 1:1:775 : 1675 : 1°75 : 1°75 : 1°725 with an average of 1 : 1°7443. Again, in fig. 2, the ratios are :— 1:38 > B°775 : 38 : 3825 with an average of 1 : 3°8. Calling the rate of the pulse z, and the number of times the first part is contained in the whole beat y, ey equals the number of times that the first part is contained in a minute, and ~ equals the part of a a minute occupied by the first part of each pulse-beat. From several observations, it was found that zy increases with 2, not directly as it, but as its eube root, consequently the following equation finds «xy in terms of 2, ay=k Sa, k being a constant, equal to 47 (about). For instance in fig. 1, ¢ = 137, y = 1°7443; | and in fig. 2, o= 44, ¥=3°8; and 137 x 1:7443 = 238-9691, 44 x 3°38 = 167°2; and 238°9691 : 167:°2 :: 1:43 : 1, and ¥1387: Y44:: == 5'155 : 3°54: : 1:456: I, which shows that in these individual cases xy varies, within the limits of experimental error, as the cube root of «. If this statement of the ratio of the first part of the trace to the whole beat is a correct one, a knowledge of the rapidity of the pulse alone is sufficient to enable the length of the first part to be found by multiplying the cube root of the rapidity by the constant quantity 47. Thus, supposing the pulse beats 64 times in a minute, the cube root of 64 being 4, 4 x 47 = 188, and the length of the first part of the beat ought to be zi, of a minute. In one case with « = 64, wy was found to be 185°75, and in another with « = 63°5, ay = 181°77, both numbers which agree closely with the requirements of the equa- tion, With x = 140, and therefore */z = 5-2, 52 x A7 = 244-4; ON THE SPHYGMOGRAPH TRACE. 17 and therefore the first part = 34; — zis PS in a pulse of that rapidity zy was found = 242°9. To save the trouble of extracting the cube root for any rapidity, these facts have been thrown into a co-ordinate form in the accom- panying table, and the observations on which the formula is based are represented by dots on their proper co-ordinates, the calculated curve, with k = 47, being represented by a continuous line. _ Since the above equation was worked out, a great many other observations have been made, several of which are recorded on the Page 354. table, and in health no cases have been found which depart from the curve more than those indicated on it. The observations made on the author are represented by simple black dots, those made on others are encircled by a ring; great size of a dot indicates that more than one independent observation has pro- duced exactly similar results. ‘In none of the cases have measurements been made after violent exercise. Differences in the height and age of the subjects experi- mented on have not been found to produee any appreciable effect. The trace from infants has not been examined. - From the equation zy = */z.k the length of the second paki of the pulse trace may be represented in terms of 2, as htop t as from the nature of y it cannot be less than unity (no pulse having been seen with two contractions or more between two successive closures of the aortic valve), the limit of cardiac rapidity may be de- duced to be 322 in a minute (k = 47); but itis scarcely probable that pulses of such a rate could remain so sufficiently long te be counted. — Tn many cases of disease implicating the circulatory system, the equation given above indicates that the duration of the first part of the heart’s action is not normal; thus, in a boy suffering from typhoid fever, on the second day after the pyrexia had ceased, and when the temperature was below the normal, zy was found = 225-25, where -# = 60, which differs from the equation 8/67 x 47 = 190°82, which shows that the length of the first part is considerably too short in the former. In the same case, three days later, the patient rapidly improving, with z = 56°5, vy = 188, which is much nearer the calculated normal result, 180°5, than on . the former occasion, the trace keeping pace with the other physical _ changes. It is probable that many other imperfections in the circulatory system can be similarly indicated, and it has been shown above with what facility a diagnosis may be arrived at. T wae c Page 17. Page 18. 18 CARDIOGRAPH TRACINGS 4. ON CARDIOGRAPH TRACINGS FROM THE HUMAN CHEST-WALL.* (Pl. II.) On applying the hand over the left pectoral region the movements of the heart can be felt with facility, especially at the end of expiration. In the following paper an attempt is made to classify and partly explain these movements, as they are reproduced by the sphygmo- graph. The earliest and perhaps the only published observations on these curves are by Dr. Marey of Paris,t who gives one trace from the human subject and others from the horse, which latter have the ad- vantage of being associated with synchronous traces from the interior of the ventricle and of the auricle. No previous observations can be found as to the relative duration of the different elements of and the other peculiarities in the human heart apex traces at different rapidities of pulse. While the subject is sitting or standing the sphygmograph can be made to give a very perfect record of the heart’s movements, as they are transmitted to the intercostal tissues, by holding the instrument horizontally with the watchwork to the right hand, the plane of the’ recording paper, and consequently of the whole instrument, being parallel to the floor, and the lever-pad at or near the point of maxi- mum pulsation, between the fifth and sixth ribs. While lying, the instrument must be held upright, as when wrist traces are taken. The movements of respiration cause so much irregularity in these traces, that it is advisable to stop breathing while they are being taken ; and this should be done at or near the end of a normal expira- tion ; it is then found that little or no effect is produced on the heart’s action, during the short time, about seven seconds, that the instrument is applied. It will also be found that quick pulses are more easily taken than slow ones, because the heart can only be rendered slow by means that make the skin cold and inelastic, or by positions that make the application of the apparatus more difficult. In all cases the spring carrying the pad should be screwed down so as to give its greatest pressure. In the account of the traces thus obtained, the rapid beats will be * “Journal of Anatomy and Physiology,” V. pp. 17-27. November, 1870. + “Physiologie Médicale de la Circulation du Sang.” Paris, 1863. Pp. 68 and 121, and elsewhere. FROM THE HUMAN CHEST-WALL. 19 first described ; after these the slow ones, by which means an idea can be best formed of the relation between curves at first sight so dif- ferent as those produced when the heart’s action is over 100 and those when it is below 50 in a minute (compare Figs. I and VI). There is a great similarity in traces from pulses above 105 to those over 140 in a minute; and the description of one will include them all. Figure I is from a heart beating 125, and it represents all the characteristic features. The movements of the lever are very extensive and sudden, so as to give the impression that they depend ‘ more on its momentum than on the heart’s action; but that such is & not the case is shown by applying the instrument a little way from Sy the point of greatest pulsation, when (as in Fig. IT) all the same * elements appear, though much less ample and otherwise modified. The main ascent commences abruptly immediately after a slight rise and fall (a, Figs. I, Il), and is always broken about midway (6) by a small fall; it is followed by a most considerable and rapid descent, which carries the lever in an unbroken line, down to a point almost as low as that from which it started. Subsequently to this comes a less sudden rise (f), which reaches about as high as the break in the main ascent; its summit is not nearly so sharp as the previous one, and from it a fall, frequently a little irregular, commences slowly, becoming more rapid, though it is interrupted by a slight rise (q), after which it continues to sink until it reaches the lowest point of the trace, from which it makes a sudden slight ascent (*), which soon becomes more gradual, continuing until the rise (a) from which the description commenced. Neglecting for the present small differences in the relative dura- tions of these components, the pulse of 140 differs from that of 110 a minute in the movements being more extensive and consequently the angles more sharp, the intermediate rates being intermediate in character. In the pulse of about 90 a minute (Fig. III) another small rise Page 19. and fall appears (¢) in addition to those previously described, which is very constant, and becomes more considerable when the heart’s actio is slower. Here also, as shown in Fig. IV in a trace taken directly after Fig. III between the sixth and seventh ribs, there is sometimes seen a reduplication of the first part of the main rise (b, c, Fig. IV), which is disguised in Fig. III probably by the momentum of the lever. Another point in which it differs from the quicker pulses is in the formation of a second undulation (b) in the main descent before it reaches its lowest point. The main ascent also is not so extensive. When about 70 beats are made in a minute, the main rise can fre- quently be shown to be doubly broken (5, c, Fig. V); but these often c2 Page 20. 20 CARDIOGRAPH TRACINGS get merged into one curved line. The subsequent fall (d) is here seen to have become much diminished, and the next rise and fall (e) of greater duration. In the slow pulses (Figs. VI, [X) the fall after the small rise pre- ceding the main ascent (a) is inconsiderable, or nil, which makes that rise appear as part of the main one, which is not the case. The rise c has now become more marked, while d has diminished so much that it is no longer the highest point of the trace, that now being at the end of the rise preceding the main descent (f), which is frequently found to be double. It is to be noticed that as the pulse gets slower, the generally ascending line between & and a gets longer; also that at all rates there is a great similarity in shape in the fall and rise between the points and &, which is quite characteristic of that part of the curve. A precise knowledge of the causes of these various changes in the direction of the human apex trace will always be somewhat deficient, from the impossibility of vivisectional verification, and from the fact that the relation of the organs concerned is different in man to what it is in animals, from which, otherwise, arguments from homology might have been more extensively employed. By means of synchronous traces from the exterior and interior of the heart of the horse Marey explains his apex trace, which in the main resembles that from the human subject. He shows that the rise, here called a, results from the contraction of the auricles, which makes it clear that that event occurs much nearer to the commencing ven- tricular contraction, represented by the origin of the main ascent, than is supposed by many. He also shows that the semilunar valves close at the break, single in his trace, in the main descent. The irre- gularities in the systolic interval he considers due to vibration of the blood caused by the tightening of the auriculo-ventricular valves, but his results were recorded after having been communicated to india- rubber tubes filled with air, and the undulations probably originating in them. It is necessary in attempting to explain these traces, especially when comparing different rapidities of pulse, always to bear in mind Marey’s most important law, that “the arterial tension (blood poten- tial) varies inversely as the rate of the pulse.”” This law, though dis- puted by some, must closely approximate to the truth, because by it so many facts with regard to the circulation of the blood are perfectly explained, that cannot be in the least accounted for elsewise; and it will shortly appear how auch it assists in interpreting the curves under consideration. After the auricles have contracted at a, the commencing ventri- cular action originates the main rise, which continues uninterrupted FROM THE HUMAN CHEST-WALL. 21- until the closure of the mitral and tricuspid valves at b. From this: point, until the opening of the aortic and pulmonary yalves, the heart’s force is expended in raising the potential of its contained blood to that of the large arteries, and it is-a well-known fact that during that time the form of the ventricles becomes somewhat globular, their diameter increasing and thus causing them to recede in their conical © pericardial cavity, producing the fall b in the quick pulses. Immediately the semilunar valves get opened, the ventricles dis- tend the proximal parts of the large arteries, and it is evident, from a what has been said above with regard to the relation of blood potential * and rapidity, that the quicker the pulse the more relaxed is the aorta at the moment before the semilunar valves open; consequently, the more rapid the pulse the greater is the disturbance of equilibrium when they do so; and as the aorta gets stretched and lengthened by Page 21. the sudden repletion, so it sends the heart forward at that moment, causing the rise d, which must therefore be greater as the pulse is quicker, which is the case. In very slow pulses the blood potential being high, the repletion of the already greatly distended arteries does little in further filling them, but acts by sending the whole mass of blood forward ; consequently the rise d is inconsiderable. The rise ¢, if not resulting from the shock of closure of the auriculo-ventricular valves, must remain unexplained. The repletion of the proximal arteries is very rapid; and the accompanying rise is overcome in quick pulses by the speedy retreat of the apex, resulting from the emptying of the heart, causing the fall d, at the end of which the ventricles cease contracting. In slow pulses the heart’s systolé is prolonged, causing slight irre- gularities in the upward tending trace, which are fairly constant (e, which is frequently double). There is evidently an appreciable interval between the end of the ventricular systulé and the closure of the semilunar valves, during which the retrograde blood-current is arriving at sufficient velocity to enable them to act; but if Marey’s law of the relation of blood poten- tial and rapidity of heart’s action is correct, we are justified in going much further, and saying that the quicker the pulse, the more slowly do the aortic valves close; for the greater the blood potential, the sooner does the heartward current become sufficiently rapid to close the valves, whose hydrodynamical relations are not otherwise modified by the rate. From these considerations, combined with the fact, in quick pulses, that just before the rise f originates the trace loses its jerky character, it is most probable that the ventricles cease contracting just before the commencing rise f (at the end of the fall d in Figs. I, II), and that the whole of the time occupied by the rise and fall f is employed in Page 22. Page 23. 22 CARDIOGRAPH TRACINGS generating the retrograde current to close the valve, the change in direction of the curve being produced, first by the relaxation of the heart causing it to advance, and then by the partial collapse of the aorta causing it again to retire. This rise and fall is also clearly shown in Fig. II, f. It can be seen in Fig. IV that in the pulse of 90 the undulation f is not so long as in the quicker ones, and in the slow curves it is shorter still, but the want of sharpness and the blending of the neigh- bouring rises prevent any accuracy being attainable in the latter cases. The immediate effect of the closure of the semilunar valves, at the end of the fall f, is to cause a check to the descent of the lever (4), as the aorta is no longer emptying itself heartwards; but this is very soon counteracted by the consequent repletion of the coronary arte- ries,* which, as can be easily shown on the post-mortem heart, in- creases the diameter of the ventricles and makes them recede, drawing the apex back, further than during any other part of the revolution. During the rest of diastolé, other minor forces come into play which are not easy to trace. Figures VII, VIII, 1X are given to show that under different con- ditions the various rises and falls may be made to assume different degrees of importance. In Fig. VIII, where the greater part of the weight of the heart rests against the chest-wall, it is particularly to be noticed that the fall after g, before which the semilunar valves close, commences from the very top of the trace, showing that the main force by which the heart is made to recede, which from the great length of the down stroke must be considerable, does not commence until after the closure of the aortic valve, which supports the theory of the cause of the active ventricular diastolé noticed above. A superficial examination of cardiograph tracings is sufficient to convince the observer that when the heart beats slowly the jirst part of the revolution, namely, from the commencing systolé until the closure of the semilunar valves, bears a smaller ratio to the whole than in quick pulses. This led the author to make a series of measurements of these ratios, on the assumption that the ventricles commence to contract at the origin of the main rise, and that the semilunar valves close at the end of the fall f. To ensure accuracy, the trace was placed on a flat piece of wood, to which was attached a ledge, along which it could be made to slide. * That the active diastolé of the ventricles results from the congestion of the coronary vessels was discovered by Briicke; and I regret that my ignorance of his observations, published in the “Sitzungsberichte der Wiener Akad. der Wiss.,” Nov., 1854, XIV. p. 845, prevented my referring to them, in a paper on the same subject, in this Journal for May, 1869. (Supra, p. 3.) FROM THE HUMAN CHEST-WALL. 23 A lever with a steel point was also in connection with the instrument, in such a way that when the tracing rested on the ledge the steel point produced scratches on the paper similar to those produced by the sphygmograph pen. _ By this means, the parts of the curve under consideration can be all projected on to one straight line, and their relative lengths measured with facility. Fig. X is a trace so prepared for measuring, and this arrangement is necessary on account of the sphygmograph lever moving in part of a circle instead of quite vertically. Further, to diminish inaccuracies in the watchwork movement, all the pulsa- tions on a trace were measured, and their average taken as the result. It was soon found that, with a given rapidity of pulse, the ratio of the first part of the heart’s revolution to the whole did not vary appreciably when traces were taken im any given position, but that when standing or sitting the first part was longer than when lying. Further, on comparing traces of different rapidities, it was found that the length of the first part varied very definitely, inversely as the rate; not so quickly, but as its square root: and the number of measurements that have been made seems to justify the law, that in health, the length of the first part of the heart’s beat varies, for a given position of the subject, inversely as the square root of the rapidity. This result differs from that of Donders,* who found that the length of the first part did not vary with different rates of heart’s action; but his means were much less efficient, he having to depend on the registration by the hand of the first and second cardiac sounds. All the facts on which the above law is supported are given in the accompanying table, in which they are thrown into the co-ordinate form, one co-ordinate, z, representing the rate of pulse, and the other, y, expressing the number of times the first part of the revolution is contained in the whole. The observations made while lying are repre- Pago 24. sented by a cross ( X ), when semi-recumbent these are encircled (@). The encircled dots (©) indicate that the position was sitting, and the standing ones are erect encircled crosses (). The simple dots are either from sitting or standing observations, but it is not certain which, as note was not taken at the time. For example, the measurement of two heart traces, one at 41 a minute while lying, and another at 141 when semi-recumbent, gave in the former case the ratio of the first part to the whole revolution 1 : 34125, in the latter, 1 : 1-832; the length of the first part is found by multiplying the rate into the number of times the first part is * “On the Rhythm of the Sounds of the Heart.” By F.C. Donders. 1865. Translated into the “ Dublin Quarterly Journal of Medical Science,” Feb., 1868. CARDIOGRAPH TRACINGS FROM THE HUMAN CHEST-WALL. 25 : éoutained in the whole, or the z by the y, when 2 is the required zy result. Thus 41 x 34125 = 139-9125, say 140; 141 x 1832 = 258312; and /41 = 6°4, about; /141 = 11°88, about; and : 6-4: 11°88 :: 140 : 259-9, about; which is very near the required numbers. If zy varies as the square root of z this can be stated thus ay = k./z; k being a constant for any given position. Taking one of the above cases, z equalling 41, zy = 140; consequently k = 22 nearly, which includes, within the limits of experimental error, all the measure- ments that have been made of traces taken while lying. When sitting or standing the results are not quite so uniform, as may be seen in the table, on which the equation curves have been drawn with k = 22, and also = 20: the latter, the lower one, passes through or approaches . most of the sitting and standing observations. To find the duration of the ventricular systolé is very important, but not at all easy in many cases. In all rapid pulses, measuring from the commencement of the main ascent to the point from which the rise f originates, and which can scarcely be anything but the termina- tion of the systolé, it has been found that that interval is contained 3°4 times in each revolution. In Fig. IV the trace of 87°5 a minute (rare from being very detailed), it is found that between the same points the ratio tothe whole is 1 : 33965, which is very near the former result. In slower pulses it is not easy to find an origin for the rise f, but if, as must be the case when the pulse is at 40 (its limit of slow- ness), the arterial tension is at its maximum, the rate of closure of the aortic valve must be at its maximum also, and the whole first part almost entirely occupied by the true systolé. When such is the case the equation zy = k./z with k = 22 is satisfied by y being equal to 3°46 (about), which is curiously near the relation found in quick pulses, and tends strongly to show, though these are all the grounds for it, that the length of the systolé of the heart is always a definite part (23th) of the whole pulsation, whatever its rate. The traces from which the preceding observations have been made were all taken on myself, and the repetition of them by others at different rates of pulse would be a means of verifying or a cause for rejecting the results arrived at. The chief sources of error in finding the ratios given above lie in the watchwork, which, if not going at an exactly similar rate each time it runs, gives the rapidity of the heart incorrectly. Also, on starting, its speed augments for a short time and then decreases, both which cause variations from the true results. By taking a trace after having remained some time in the hot room Page 25. Page 27. 26 CARDIOGRAPH TRACINGS FROM THE HUMAN CHEST-WALL. of a Turkish bath very rapid pulses can be recorded up to and above 150 a minute in health, without the least inconvenience. Very slow pulses can be produced by lying nude some time in a cold air, or by drinking iced water, especially when nude. _ DESCRIPTION OF THE FIGURES. ’ They were all taken one-half the size here represented. Except Fig. IX, they read from left to right. Fig. I. Apex trace of a heart beating 125 times a minute. Fig. If. Trace of a heart beating 125 a minute, half an inch prays. cat apex beat. Fig. III. Apex trace of a heart beating 88 a minute. * Fig. IV. Trace between 6th and 7th ribs, below the apex, of a heart beating 88 a minute. Fig. V. Apex trace of a heart beating 76 a minute. Fig. VI. Apex trace of a heart beating 50 a minute. Fig. VII. Apex trace of a pulse of 90 a minute, taken with a lever two inches long, attached to the sphygmograph pad by‘a thread. Fig. VIII. Apex trace of a pulse at 60 a minute, taken when the body was inclined forward. Fig. 1X. Apex trace of a pulse at 42 a minute. It reads from right to left. Fig. X. Apex trace of a pulse at 103 a minute, prepared for measuring. CONSTRUCTION AND USE OF A CARDIO-SPHYGMOGRAPH. 27 5. ON THE CONSTRUCTION AND USE OF A SIMPLE Page 265. CARDIO-SPHYGMOGRAPH.* Ir is evident that a precise knowledge of the intervals between the main elements of the cardiograph and the sphygmograph trace must be of value in studying the hydrodynamics of the circulation of the blood; and a description will be here given of an instrument by which several results of interest have been obtained on this subject. This cardio-sphygmograph consists of a piece of board, 10 inches long by 54 inches broad, and is about half an inch thick, along one side of which one of Marey’s sphygmographs can be fixed, as shown in the accompanying figure. On the opposite to this is a spring (a), similar to that employed in the sphygmograph, which is attached toa movable support (b), so that its strength may be modified. A small ivory pad (c) is fixed to the lower surface of the free end of the spring, and this isin communication with the recording lever of the cardiograph apparatus by means of a silk thread (d). In this instrument the cardiograph lever (e) is very light, a little over two inches long, and connected to the board by means of a frame (/), which is just free of the movable part of the sphygmograph, when that is in position. The lever, which is one of the third system, is connected on either side, close to its fixed end, to two silk threads, one of which (d) is attached to the cardiograph spring, and the other to a small spring (g), which Page 266. moves it when it is less acted on by the stronger one. The apparatus is so arranged that the lever works perfectly when it is so placed as to be above the recording paper of the sphygmograph, when the latter is in position. The tip of the lever carries a steel pen (k). * “Journal of Anatomy and Physiology,” V. pp.-265-70. May, 1871. Page 267. 28 CONSTRUCTION AND USE OF A The apparatus therefore consists of a cardiograph and of a sphyg- mograph, and these are so fixed that they both record on the same paper ; and the object to be attained is to get them both to record at the same time, the one the movements of the heart’s apex, the other the dilatations of the artery at the wrist. To obtain this result the sphygmograph is first fixed, as usual, on the left arm, and the recording paper is adjusted to its place on the watchwork, With the cardiograph in the right hand, the left arm is then moved until the attached instrument rests on the board in the position shown in the figure, and when there, it is maintained in its place by certain pegs and holes in the board, which respectively come in contact with the main parts and receive the projections of the instru-. ment. The arm and attached instruments are then moved until the pad of the cardiograph spring is brought in contact with the spot, generally between the 5th and 6th ribs, at which the heart’s pulsations are most marked; the position of the pad in relation to the board having been previously sd fixed as to enable this to be done with facility, the whole being maintained in the horizontal position. The contact of the cardiograph pad with the chest-wall causes the lever to recede from the chest, and it is allowed to do so until its pen arrives above the recording paper; the whole apparatus being steadied by the right hand. When the levers of the two instruments are both found to be moving freely, the watchwork of the sphygmograph is set in action by a string (m) held by the right hand, and at the other end connected with the stop-block of the train of wheels. The two levers recording on the smoked paper give a combined trace of which Fig. II is an example. As with simple cardiograph traces it is advisable and almost neces- sary to hold the breath while the trace is being taken, and further, to simplify the working of the instrument, the chest should be empty at the time. It is evident from the above description that the two levers write in opposite directions, and consequently this figure must be turned the other way up that the cardiograph trace may be properly. seen, and then it must be read from right to left, not from left to right, as the sphygmograph trace. SIMPLE CARDIO-SPHYGMOGRAPH. —~ 29. The commencement of the two traces is indicated by the curved lines to the left of each trace as they are looked at without moving the page, and these curved lines are produced by letting the levers move without the watchwork, whilst the instrument is being fixed in posi- tion. Synchronous points in the two traces must evidently be at equal distances from the starting points in the traces, and therefore the one can be projected on the other by compasses or by super- position. 3 In all cases it is necessary, both in the cardiograph and in the sphygmograph trace, to project all the main points, such as the origin of the main rise, and the deepest point in the seeondary fall, on to one line in the trace ; for, as the levers move in part of a circle, any point at the summit of the trace, if projected straight downwards, would not be correctly related to the lower parts of the trace. This correc- tion is best made by a most simple arrangement; a flat piece of board has a straight slip of wood fixed close to one edge; against this the tracing rests, being supported on the board. Two nails are fixed on the board, so that they bear the same relations to its supported trace as that did to the axes of the levers which marked on it in the cardio- sphygmograph. The marking apparatus consists of two pieces of string, each fixed at one end to the nails, and at the other carrying needles ; these pieces of string must be of the same length as the _ levers to which they correspond, the points of the needles must pierce them, and the other ends of the needles must be attached to the nails by a thread to prevent them from moving irregularly. The cardio-sphygmograph can be best applied when the person Page 268. using it is sitting, as it can then be made to rest on the arm of a chair, and in practice it is better not to have the main part of the instru- ment press against the chest-wall, as if it does the heart’s movement imparts itself to the whole apparatus, and so complicates the trace. In considering the results arrived at by the use of this instrument, it will be necessary to define a few of the terms that have to be employed in explaining them. (1.) The first cardiac interval is that which occurs between the commencing systolé and the closure of the aortic valve at the heart. (2.) The first arterial interval is that which occurs between the indications of the commencing systolé and the closure of the aortic valve in an artery. The radial artery at the wrist is the only one that is here considered. As the commencement of the arterial rise is somewhat later than the commencing systolé at the heart, and as the difference between the first cardiac interval and the first arterial interval is not great, these two events coincide in part of their duration, and give rise to minor divisions, which may be thus named and defined. ale ai Page 269. 30 CONSTRUCTION AND USE OF A (3.) The first cardio-arterial interval is that which occurs between the commencing systolé at the heart and its indication in an artery (the radial). (4.) The conjugate cardio-arterial interval is that which occurs between the commencing systolic rise in on artery and the closure of the aortic valve at the heart. (5.) The second cardio-arterial interval is that which occurs between the closure of the aortic valve at the heart and its indication in an artery.* On comparing the lengths of the first cardio-arterial interval with different rates of pulse, it is found that as the pulse is slower, so this interval is longer, and that its length does not increase as rapidly as the pulse beat, but as its square root. Consequently if the number of times that the first cardio-arterial interval is contained in its component pulsation is represented by z and the rapidity of the pulse by x, then ke = k./w, and measurements show that the constant quantity & equals 39 (or perhaps 39°25) for the sitting posture. A knowledge of this equation, therefore, gives a means of calculating the length of the first cardio-arterial interval when the rapidity of the pulse is known; and as the first cardiac interval also varies as the square root of the pulse beat,} it is evident that from its definition the first cardio-arte- rial interval must be a constant part of the first cardiac interval, whatever the rate, and this has been found to be the case by inde- pendent measurement. Another necessary result from these equations is, that the conjugate cardio-arterial interval varies inversely as the square rovt of the heart’s rapidity. The length of the second cardio-arterial interval can also be found by subtracting the length of the conjugate cardio-arterial interval from that of the first arterial interval, which varies inversely as the cube root of the pulse rate,t and by this means it has been found to vary very little with different rapidities of pulse, being a little longer in the slower pulses. In the sphymograph traces of slow pulses the major descent of the first arterial interval is broken by a notch, and it is found that the * In the above definitions it has been assumed that the sphygmograph trace gives indications of the closure of the aortic valves ; and in the measurements to be referred: to below, the secondary rise, which puts so abrupt a termination to the major fall in each pulsation, is considered to be caused by the closure of these valves, as generally assumed ; though Dr. Sanderson has arrived at a different conclusion (“Medical Times and Gazette,” March 25, 1871), from evidence which seems to be anything but convincing. + “Journal of Anatomy and Physiology,” Nov., 1870. “ On Cardiograph Traces from the human chest-wall.” (Supra, p. 18.) t “ Proceedings of the Royal Society,” No. 120, 1870. “On the relative duration of the component parts of the radial sphygmograph tracein health.” (Supra, p. 14.) SIMPLE CARDIO-SPHYGMOGRAPH. 3l deepest point of this notch is always exactly synchronous with the point of closure of the aortic valve. This leads to the almost neces- sary conclusion, that the subsequent slight rise or change in direction of the trace is the result of the simultaneous movement of the whole column of blood produced by the shock of the closure of the aortic valve; the secondary rise at the commencement of the second arterial interval being the more slowly transmitted pressure wave, which started at the same time This explanation being correct, it is evident that the results obtained, by measuring the number of times that the interval between the origin of the main arterial rise and the bottom of the notch in the major fall is contained in the first arterial interval, ought to give the same results as those obtained by dividing Page 270. the calculated length of the conjugate cardio-arterial interval into the first arterialinterval. Such has been found to be the case very closely, but a sphygmograph trace must be a very good one to show the notch in the first arterial interval sharply defined, and the subsequent rise commencing abruptly. In all the cases above discussed it has been assumed that the subject from whom the traces were taken was sitting at the time, and as the length of the first cardiac interval changes for the same rate of pulse with change of position, it is evident that the equation given above (namely, zz = 39,/z) must be changed also; and the change probably consists in increasing the value of the constant for the stand- ing posture, as then the first cardiac interval is shorter, but still varies inversely as the square root of the heart’s rate. The explanation given above of the cause of the notch in the first arterial interval, might lead to the expectation that the commencing ’ cardiac systolé indicates itself at the wrist in the same way ; but there is no such marked change of direction in sphygmograph traces, though a slight rise is generally seen just before the main ascent originates, especially in pulse of about 70 in a minute; and it is not improbable that the second rise in the extremely dicrotic pulse of adynamic pyrexia is caused by a combination of the slow pressure wave resulting from the closure of the aortic valve, and the sudden onward motion given to the whole mass of blood in the vessels at the moment of opening of the aortic valve at the commencement of the next systolé. The facts on which the above equations have been based are pub- “a lished in the “‘ Proceedings of the Royal Society,” XIX. No. 126, p-318.* “4 - Mr. Hawksley, of Blenheim Street, is constructing a cardio- sphygmograph from the model described above, with a few minor improvements, which can easily be applied in the study of pathological conditions. * Infra, p. 32. Page 318. Page 320. © 32 MUTUAL RELATIONS OF THE APEX CARDIOGRAPH 6.ON THE MUTUAL RELATIONS OF THE APEX CAR- DIOGRAPH AND RADIAL SPHYGMOGRAPH TRACE.* A vrstre to acquire an accurate knowledge of the relation born by the commencing contraction of the heart to the origin of the primary rise in the pulse at the wrist, led the author to construct an instrument which has enabled him to determine, with considerable accuracy, the mutual relation of these two points, and to demonstrate one or two unexpected results, not altogether without interest. [Here follows an account of the cardio-sphygmograph described in the preceding paper.—Ep. | . All the observations were made on the same subject, aged 24, in good health. They were all made in the sitting posture, as the appa- ratus could then be held more firmly, or rested on the arm of a chair. To facilitate description, the following terms and symbols will be employed with regard to pulse-traces. 1. The rapidity of the pulse is symbolically represented by z. 2. The first cardiac interval is that which oceurs between the com- mencement of the systolic rise and the point of closure of the aortic valve, in cardiograph traces. The number of times that this interval is contained in its component beat is represented by y; and the law as to its length, published spatiale eb will be assumed. It may be stated thus :-— vy = 20/2. 8. The first arterial interval is that which occurs between the com- mencement of the primary rise and the termination of the major fall in arterial sphygmograph traces. The number of times that this interval is contained in its component beat is represented by y'; and the law as to its length at the radial artery, which is alone considered in this communication, published in the “ Proceedings of the Royal Society ” (No. 120, 1870),t will be assumed ; it may be thus stated :— ay’ = 47 8/2. 4. The first cardio-arterial interval is that which occurs between the commencement of the systolic rise in the cardiograph trace and the origin of the main rise in the sphygmograph trace. The number of times that this interval is contained in its component beat is repre- sented by z. 5. The conjugate cardio-arterial interval is that portion of the first * “Proceedings of the Royal Society,” XIX. pp. 318-24. Read Feb. 23, 1871. + “ Journal of Anatomy and Physiology,” Cambridge, V. Nov. 1870. (Supra, p. 18.) t Supra, p. 14. ’ AND RADIAL SPHYGMOGRAPH TRACE. 33 cardiac interval which is synchronous with a portion of the first arte- rial interval. It is therefore the interval between the commencing sphygmograph rise and the point of closure of the aortic valve as repre- sented in the cardiograph trace. 6. The second cardio-arterial interval is that which occurs between the point of closure of the aortic valve and its indication at the artery under consideration. In commencing to work with the cardio-sphygmograph, measure- ments were made to find the duration of the first cardio-arterial interval, as it required but a few experiments to prove that the heart commences to contract before the pulse is indicated at the wrist. By means of compasses, or by superposing one trace on the other, the commencing cardiograph rise was projected on the sphygmograph trace ; and the interval between this event and the origin of the radial rise was then measured into its component beat in each pulsation of Page 321. the trace, from which the average of the observation was obtained. The results are given in Table I, Column I; and in Column III some of these are expressed in parts of a minute, whereby a better idea can be obtained as to their significance. Taste I. wl nly B 4 R : i 002768 “002802 -002524 “002538 002222 “002229 00197 “003 957 SERSSeReesRAseS Ro 6 60 9 hh ei ie on on & Pas” apeaeeea" » oH From these results itis seen that the first cardio-arterial interval is longer in slow than in quick pulses, and that it does not increase as quickly as the pulse diminishes in rapidity; but that the statement that it varies inversely as the square root of the rapidity is correct, or very nearly so, is rendered evident by comparing Columns III and IV, in the latter of which the duration of the first cardio-arterial interval D Page 322. 34 MUTUAL RELATIONS OF THE APEX CARDIOGRAPH is calculated from the formula #z = 39 /z. The chief source of error in these observations is the slight uncertainty in the rate of movement: of the watchwork of the instrument, on which the calculation of the rapidity of the pulse depended. a On comparing this equation, namely wz = 39,/2, with the one above referred to as to the relations of the first cardiac interval, namely, ay = 20/z, it is evident that the length of the first cardio-arterial interval is ‘5128, or just over half that of the first cardiac interval, whatever the rate of the pulse. : This being the case, a more precise method is acquired of verifying the results arrived at; for by finding the number of times that the first cardio-arterial interval is contained inthe first cardiac interval, a constant quantity ought to be the result, which is independent of the rapidity of the pulse. Table II contains these measurements; and it may be seen that, though there is a small range of variation, the num- bers are all very near to the theoretical requirement, which is 1:95; and their average is 1°983. Tas.e II. { Sak cesta 2 Number of times ii that the first ae that the first Rapidity | cardio-arterial | P®PIdity | cardio-arterial of interval of interval is contained is contained pulse. in the first car- pulse. in the first car- diac interval. diac interval. 58 1°95 81°5 1°95 64 2° 83 2:1 69 1°9 84 2-1 70 1°9125 85 1°995 71 1:975 85°5 1°95 72 2-058 86 2° 74, 1°975 88°5 2°05 76 1°98 91 2°15 78 1°9 92 1°85 79 1°925 94, 1°95 79°5 1°9 97 2°15 80 1°95 154 1°975 It is generally known that in the sphygmograph traces of most slow pulses there is a notch in the first arterial interval, immediately pre- ceding the major fall; and one of the most marked results of the use of the cardio-sphygmograph is the determination of the fact that the point of closure of the aortic valve at the heart is always exactly synchronous with the lowest part of this notch, or the point of abrupt change of direction in the major fall of the sphygmograph trace. This leads to the almost necessary conclusion that the subsequent slight rise cr change in direction of the trace is the result of the simul- AND RADIAL SPHYGMOGRAPH TRACE. 35 taneous movement of the whole column of blood produced by the suddenness of the shock of closure of the aortic valve, the secondary rise in the same trace being the more slowly transmitted pressure wave resulting from the same cause. The slower the pulse the more distinct is this notch ; and by com- paring different rapidities, a gradual diminution in its conspicuousness is apparent, it rising higher and higher above the point of termination of the major fall as the pulse is quicker and quicker. When the heart’s rate is about 75 in a minute, the notch is halfway down the major descent, and is partially blended with it; when over 100 a minute, as the aortic valve cluses when the ascent is at its maximum, the notch is so blended with the pressure wave as not to indicate itself separately. In slow pulses, the systolic main rise being quite over when the aortic valve closes, the shock wave indicates itself by an abrupt but Page 323 not considerable rise, breaking the very gradual major descent. This explanation being correct, another means is obtained of check- ing the results arrived at by the combined instrument; and Table III, Column IT, contains a few measurements of the number of times that the conjugate arterial interval is contained in the first arterial interval, _ as found by measuring the ratio of the interval between the commenc- ing arterial rise and the bottom of the notch in the major fall to the whole first arterial interval. Column III gives the theoretical results necessitated by the equations given above. Taste III. Number of times the conjugate cardio-arterial interval is Rapidity contained ra the first of — as found from as calculated measurement (approxi- of mately). radial trace 37 1°595 1°6 45 1-635 1 “625 58 1°69 59 1-7083 1-72 60 1°74 1-734 68 1-797 1°78 It may be mentioned that the reason why so few of these instances are given, is, that there is considerable difficulty in measuring these D2 36 MUTUAL RELATIONS OF THE APEX CARDIOGRAPH. small intervals into one another with precision; but by practice a very fair estimate can be made of their value, and in all cases they seem to agree with theoretical requirement. The close accordance of the results obtained by this method in very slow pulses, and the calculated results arrived at from facts relating only to quicker ones, tends strongly to establish the correctness of the law given with regard to them. . In Table IV the lengths, in parts of a minute, of the different intervals referred to in this communication, are given as calculated from the equations on which they have been shown to depend. With regard to the second cardio-arterial interval, a reference to Column VII will show that it varies very slightly within the range of the heart’s action, not being 4 longer in a pulse of 36 than in a pulse of 169 ina minute. Page 324. Taste IV. I. II. III. TV; V. VI. Vit. Length of Length of | Length of | second Rapi- | Length of Perper Pires oat first cardio- | conjugate | cardio- dity of pulse-beat, jbooed ats ct arterial | cardio-arte-| arterial y in parts of Peqtcne of re alt ’ € interval, in | rial interval.| interval, in pulse. | a minute. | RRA ee wis parts of | in parts of | parts ofa * | @minute. | a minute. | a minute. | minute. 36 | 027 ‘0083033 | 006428 0042735 | °0040298 | ‘00239821 49 | *020408 | :00714286 | -005813 003663 00347986 | °00233342 64 015625 "00625 005319 "003205 “0030449 00227425 81 0123457 | *005 “00491356 | -0028474 | ‘0027081 00220546 100 “01 "005 | 004.6234 *0025641 *0024359 00218745 121 | 0082645 | -0045_ “004299 002331 00221445 | +0020847 144, “00694, “00416 “004.0486 *0021365 *0020301 “0020185 169 | 005917 | -003846 ‘0038485 | -0019704 | 0018756 | ‘0019729 In conclusion, the following are the results that have been arrived at by the use of the above cardio-sphygmograph :— 1. The first cardio-arterial interval varies inversely as the square root of the pulse-rate. 2. The conjugate cardio-arterial interval varies inversely as the square root of the pulse-rate. 3. The second cardio-arterial interval varies very little with differ- ent pulse-rates, but is slightly longer in slower pulses. 4. The depth of the notch in the first arterial interval of the sphygmograph trace occurs at the moment of closure of the aortic valve. 5. There is no definite indication in the sphygmograph trace of the moment at which the cardiac systolé commences. > RELATION OF TEMPERATURE, ETC. 37 7. ON THE RELATION OF THE TEMPERATURE OF Page 126. THE AIR TO THAT OF THE BODY.* Tue nature of the evidence affecting theories in biological science is generally so far from direct, that it is only by the systematic working out of many of the necessary deductions that any idea can be formed A as to their value. The experiments detailed in this communication : were suggested by the theory to be referred to immediately, and their close agreement with its requirements tends strongly to substantiate its accuracy. In a paper published elsewheret I have detailed several observa- tions which tend to show that many of the minor fluctuations in the temperature of the human body result from alterations in the amount of blood exposed at its surface to the influence of external absorbing and conducting media. Others have repeated these experimentsf{, and obtained the same results. For example; on stripping the healthy body in an air of about 50° F., a rise of the internal temperature (judged by that of the floor of the mouth) commences immediately, and in about half an hour amounts to as much as three-fourths of a degree. According to the above-mentioned theory this phenomenon is explained thus; the con- tact.of the cold air against the surface of the skin, previously main- _ tained at a much higher temperature by the clothes covering it, pro- duces so considerable a contraction of the cutaneous muscular vessels, and the blood is driven so far inwards, that the conducting power of the thus modified skin is rendered considerably less than that of the clothes and blood-filled skin combined, in the previous condition; consequently the body temperature rises until a higher equilibrium is attained. Such being the case, and the contraction of the cutaneous vessels being evidently caused by the cold, it is more than probable that the amount of this contraction should depend on the degree of cold ap- plied, that is, on the temperature of the external air in which the observation is being conducted; and the extent of this action would Page 127. manifest itself by its effect on the body temperature, less cutaneous contraction causing less diminished conduction and consequently less rise of temperature on stripping. Se re * “ Journal of Anatomy and Physiology,” VI. pp. 126-30. Noy. 1871. ‘ “ Proceedings of the Royal Society,” No. 112, 1869, p. 419, e¢ seg. (Suprd, p. 6. t J. F. Goodhart, “ Guy’s Hospital Reports,” 1869. 38 RELATION OF THE TEMPERATURE OF THE AIR Similar reasoning would lead us to anticipate a temperature of air sufficiently high to produce exactly as much cutaneous contraction as will make up for the loss of the clothing, and consequently no change in the body temperature on stripping. The correctness of these deductions may be judged from the fol- lowing observations, which were all made under similar conditions, on myself, while standing. zB af. Time. | Temperature. Ses grtagc Time. |Temperature. ive: cn 11.15 98°95 ) 11.15 98°8 ) 11.20 11.20 11.25 98°975 11.25 98°8 11.30 98975 11.30 98°8 11.35 11.35 11.40 99:3 . azo. | 11.40 99° p bane 11.45 99°35 11.45 : 11.50 99°575 11.50 99°35 11.55 99°625 11.55 12 NIGHT} 99°7 12 NIGHT} 99°375 J 12.5 99675 |) Stripped at 11.30. Rise 0°7° F. Stripped at 11.30. Rise 0°575° F. IIT. IV. | Y Temperature 2 Temperature Time. |Temperature. Ee yt Time. |Temperature. oF Ais. 10.45 98°1 7 11 P.M. > 10.50 98° 11.5 99° 10.55 97°925 11.10 99° 11 P.M. 98: + 59° F 11.15 99° 11.5 98°19 4 11.20 99° . 67°F. 11.10 98°35 11.25 99°05 11.15 98°4 11.30 99°19 11.20 98°425 J 11.35 99°21 11.40 99°19 4 Stripped at 10.55. Rise 0°5° F. Stripped at 11.15. Rise 0°2° F, a TO THAT OF THE BODY. 39 Page 128 Ke Vi. Time. /|Temperature. sp cane Time. | Temperature. ee? react 10.15 99°15 7 10.15 98°8 >) 10.20 99°15 10.20 98°875 10.25 99°15 10.25 98°95 10.30 99°125 10.30 98°95 - 71°F 10.35 99°025 71°F 10.35 99° 10.40 99°125 pre ; 10.40 99° 10.45 99°175 10.45 99° 10.50 99°175 10.55 997175 11 Pm. 99°175 a Stripped at 10.30. Rise at 0°05° F. Stripped at 10.25. Rise 0°05° F. Vil VIII. 5 T tur J T ture Time. | Temperature. rar ee ©] Time. Temperature. an 10.30 99°21 } 10.15 98°8 ry 10.35 99°205 10.20 98°825 10.40 99°2 10.25 10.45 99°3 10.30 98°9 10.50 99°3 10.35 99° . 73°F 10.55 99°15 - 72 R 10.40 . 11 P.M. 99°35 10.45 99°1 11.5 99°3 10.50 99°1 11.10 99°325 10.55 99°71 11.15 99°4 11 P.M. 99°1 J 11.20 99425 =|) Stripped at 10.50. Rise 0°125° F. Stripped at 10.36. Rise 0-1° F. It is readily seen that the hotter the air, the less is the stripping rise, and that when an external temperature of 70° F. is reached, there is no rise at all. Several of the higher temperature observations are here given and but few of the lower, because in the paper above re- ferred to (“‘ Proceedings of the Royal Society,”’ No. 116, 1869)* there are six or seven of the latter recorded with the temperature noted in all. From these facts it may be clearly seen that in air below the tem- perature of 70° F., the stripping rise varies inversely as the temperature. Page 129. Experiments as low as to 45° F., have been made, but no limit has * Supra, p. 6. 40 RELATION OF THE TEMPERATURE OF THE AIR been reached in that direction yet. Above 70° F. of the air, the results are modified by the sweating that always accompanies so high a tem- perature if the body is clothed, and quite a different class of pheno- "mena appear, which have not been much studied. Page 130. In V and VII there was a slight fall of temperature at the moment of stripping ; this was probably connected with the perceptible moisture on the surface which evaporates almost immediately the clothes are removed. With regard to the nature of the clothing removed. It always had considerable non-conducting power, being composed in all cases of at least two layers of woollen material; though, as the observations in the warmer air were made in summer, and those in the colder, during corresponding seasons, the dress worn varied with the time of year, being thinner in the former and thicker in the latter. On the whole the amount of clothing worn does not seem to affect the results as long as there is sufficient to keep the body warm under ordinary cir- cumstances; and in the English climate to do this, one woollen cover- ing seems always essential. Some results obtained by Dr. V. Weyrich* with regard to the hygrometric condition of the skin at different temperatures of the atmosphere, obtained by means of an hygrometer specially adapted for the purpose, bear so fully on the subject under consideration that they will be here given. lst. When the body is clothed, the amount of moisture excreted by the skin does not vary appreciably when the observations are con- ducted in an air below 70°F 2nd. When above 70° F. the amount of moisture excreted by the skin rapidly increases with a rise in the temperature of the atmo- sphere. It is thus seen that by means of an entirely different method of observation, Weyrich finds that in an air of: 70°F. sweating com- mences; and by a combination of his results with those arrived at from the facts given above, the following conclusions may be drawn :— With regard to the human body, when covered with badly con- ducting clothing, 70° F. is a critical temperature of the atmosphere. The removal of the clothing at that temperature produces sufticient contraction of the cutaneous muscular arteries to counteract the cool- ing effects of its loss, and consequently the internal temperature does not change; whilst on stripping at lower temperatures, the vascular contraction induced more than makes up for the covering lost, and is * “Die Unmerkliche Wasserverdunstung der Menschlichen Haut,” Leipzig, 1862. Abstract in “ British and Foreign Medical Chirurgical Review,” Oct., 1863. TO THAT OF THE BODY. 41 consequently followed by a rise of internal body temperature; which, like the vascular contraction, is greater as the cold is more consider- able. Above 70° F. the amount of perspiration varies with the degree of heat and so far compensates, by evaporation, for the differ- ences of temperature as to maintain the body at a nearly uniform The temperature of air at which sweating begins in the nude body is not known. It is at about 86° F. when standing at rest. Page 446. 42 ON PULSE FREQUENCY, 8. ON PULSE FREQUENCY, AND THE FORCES WHICH VARY IT.* THE circulation of the blood is a uniform circulation, the pulsation being neglected, and a uniform circulation is one in which the quantity of fluid flowing through all segments of the circulating system is the same; otherwise there would be a tendency for the fluid to accumulate at certain points, which is contrary to the premises. To arrive at precise conclusions respecting the circulation there are two points which must be considered :—1st., The laws which regu- late the flow of fluids through capillary tubes. 2nd., The variations in the capacity of the circulating system under different pressures. These will be considered separately. Poiseuille found that the flow of fluids through capillary tubes varies directly as the pressure and as the fourth power of the diameter of the tubes. The author has verified the former of these results on the vessels of the animal system by a different method. Respecting the capacity of the arteries and ventricles under different blood-pressures, it is evident that the capacity of the former must depend on the pressure only, for they are simple elastic tubes, and must be more capacious under high than under low pressures; reasons are given below for a more precise state- ment of this relation. To maintain a uniform circulation with a pulsating motor, like the heart, it is evident from the above considera- tions that variations in the resistance at the small arteries must pro- duce variations in pulse-rate; and that unless the capacity of the arteries and heart vary directly as the pressure, variations in blood pressure must be also attended with change in pulse frequency. That the capacity of the ventricles is dependent on the arterial blood pres- sure can be proved by the varied amount of opening up of the ventricular cavities which follows different fluid pressures in the coronary arteries. Next, the different forces which vary the pulse-rate must be con- sidered. It can be shown that any change in the resistance to the flow of blood through the capillaries varies the pulse-rate, increased re- sistance rendering the pulse slower, and the reverse. As instances of these effects may be given, the pulse-slowing effects of stripping in a cold air, of a cold bath, and of compression of large arteries; the * “ Nature,” VI. pp. 446, 7. Sept. 26, 1872. Paper read before the British Association at the Brighton Meeting in Section D, Department of Anatomy and Physiology. See “ Report of British Association,” 1872, p. 151. AND THE FORCES WHICH VARY IT. 43 pulse-quickening effects of a hot bath, whether air or water. Nume- rous experiments by the author prove that the effect of copious blood- letting is not to modify the pulse-rate at all, thus showing that the law given by Marey respecting pulse frequency is not correctly based. The above points, namely, the law of Poiseuille, the dependence of the capacity of the arteries and ventricles on the pressure of the blood, the dependence of the pulse-rate on the peripheral resistance and its non-dependence on the blood pressure, can ull be correlated by only one theory, namely, that the heart always re-commences to beat when the tension or pressure in the arteries has fallen at invariable proportions, which also assumes that the capacity of the heart and arteries varies directly as the pressure. The facts that the arteries are generally empty after death, and that the cavity of the heart is sometimes found to be obliterated on rigor mortis, show that absence of pressure and capacity go together. This theory explains the known peculiarities in pulse-rate attending change in position, by showing that while standing all the pressure of the body weight is borne by non-compressible rigid tissues, and so the circulation is normal, but while lying, the soft parts are compressed and resistance introduced into the circulation, reducing the rapidity of tension-fall, and therefore the frequency of the pulse; an interme- diate condition attends the sitting posture. The pulse quickens during inspiration, and becomes slower during expiration; for during the former act the reducing pressure in the chest lowers the aortic blood pressure, and makes the tension-fall more rapid, while in expiration the reverse occurs. This theory also is the only one which throws light on the cardio- graph law published by the author (see “Journal of Anatomy and Physiology,” 1870-71), which may be thus stated:—For any given pulse-rate the first part of the heart’s revolution has a constant length, but it varies as the square root of the length of the complete pulsa- tion. The pulse-rate not depending on the blood pressure, and the length of the first cardiac interval not varying when the rate is con- stant, its length also does not depend on the blood pressure. The first cardiac interval may be divided into the systolé and the interval * between that and the closure of the aortic valve (the diaspasis) ; these combined not varying as the blood pressure, it is almost certain that separately they do uot do so either ; so it may be said that neither the length of the systolé nor of the diaspasis depends on the blood pressure. But the fall of tension between the pulse-beats being but small, and the diaspasis length not depending on the blood pressure, there is no reason why it should vary in length with different pulse- rates; and assuming this in connection with the measured diaspasis length in a particular case (‘00183 of a minute), it can be deducted 44 ON PULSE FREQUENOY, ETC. from the above cardiograph law, that the systolic length varies as the square root of the diastolic. From these facts the relation of the nutrition of the heart to the time of heart nutrition (diastolé) and to the blood pressure may be deduced; for the systolic length not varying with the blood pressure when the pulse-rate is constant, it is evident that the cardiac nutrition must vary directly as the blood pressure in the aorta; and the systolé varying as the square root of the diastolic time, shows that the nutrition of the heart varies as the square of the time of nutrition (diastolé), for with a quadruple resis- Page 447. tance to the peripheral circulation, the heart would be four times the time in emptying itself, but it is only double that time, which demon- strates the statement. A complete logical explanation of the action of the pneumogastric can be given on this theory, by assuming that its function consists in diminishing the calibre of the small arteries of the coronary system, and always keeping them somewhat contracted. LAW REGULATING THE FREQUENCY OF THE PULSE. 45 9. ON THE LAW WHICH REGULATES THE FRE- QUENCY OF THE PULSE.* | THE paucity of mechanical theories to explain the frequency of the pulse, probably arises from the very general assumption, that in all cases when the rapidity of the heart’s beat is caused to vary, the Page 1, action of nerves with special powers of retarding or quickening it is ~ brought into play ; and the relation of heart power to work to be per- formed, has not been introduced into the problem. The theory of energy has of late spread so far and wide the neces- sity for finding in all cases where work is done a sufficient source for the production of that work, in one form or other, that a vague state- ment to the effect that heart frequency depends solely on nerve action, is far from sufficient for the requirements of physiologists. It is now necessary to show that with different amounts of work to be per- formed in the circulation, different supplies of nutrient substance must be presented to the motor organ, just as in the steam-engine the amount of fuel must be varied according to the work required from the machine. When the microscope revealed the existence of a well-marked muscular coat to the smaller systemic arteries, it became evident that the different diameters of those vessels consequent on the degrees of contraction of their walls, varied the amount of force necessary to propel the blood through them ; and these variations have been con- -siderably studied of late. Dr. Marey of Paris, the introducer of the sphygmograph, has, in his most scientific treatise “‘ On the Circulation ‘of the Blood,”’+ strongly drawn attention to this subject, and he has worked ont a theory respecting the law regulating the frequency of the pulse, which is based mainly on the variations in arterial resist- ance. This theory of Marey’s it will be necessary to recapitulate here, and to examine the facts on which it rests. The following is the law in the two forms in which he gives it. 1. “ The heart beats so much the more frequently, as it experi- ences less difficulty in emptying itself.” * This paper was originally printed and published separately (London: H. K. Lewis, 136, Gower Street, 1872), but was afterwards republished in the “Journal of Anatomy and Physiology,” VII. pp. 219-32, and VIII. pp. 54-61, with one or two corrections which have been here inserted.—Eb. + “ Physiologie Medicale de la Circulation du Sang.” Paris, 1863. Page 2. Page 3. Page 4. 46 THE LAW WHICH REGULATES 2. “The frequency of the pulse varies inversely as the arterial tension.” As reasons for the accuracy of this law are given— Ist. The analogy of other intermittent muscular movements, as the following :—A man can walk a certain distance quicker, the less he is loaded. Or this—The hand can be moved alternately backwards and forwards more quickly in air than in the more resisting fluid, water. 2nd. The pressure can be made to change by variations in the amount of blood in circulation, and by modifications in the degree of arterial or capillary resistance, both of which vary the pulse-rate in the manner required by the theory. To prove the effects of different amounts of blood in circulation, the experiments of Hales are quoted, in which he found that loss of blood increased the frequency of the pulse. To prove the effects of varied arterial or capillary resistance, many satisfactory and original results are referred to; among them, the effect of compressing the abdominal aorta, or the femorals, which retards the pulse; the effects of cold baths, according to Drs. Bence Jones and Dickinson, when the pulse was greatly reduced in fre- quency ; the quickened pulse following successive additions of warm clothing over the body is also proved. : From these latter results it is clear that Marey assumes that by varying the capillary resistance the blood pressure is also varied at the same time, but this assumption is not necessarily true in a circu- lation that is maintained by a pulsating motor organ whose rate is variable, as can be easily shown by an analogy from electricity, which is a useful one in many ways to students of the circulation, and is quite worth being worked out by each. It is this :—Suppose a battery connected, through a break-and-make key, to a long uniform insulated line or telegraph cable, insulated at the other end, and connected with a static galvanometer. First connect the two parts by the key and thereby charge the line, and then break connection; upon this the charge will fall in tension slowly, and this fall may be observed on the galvanometer; when the tension has fallen one half, reconnect and break again. It is evident that if this process be repeated, a definite current is maintained between the cable and the surrounding bodies to which it leaks. If the line be now halved in length, whereby the resistance is doubled, and again insulated at the free end, it is evident that by again break- ing and making contact as before, when the tension is halved, the maximum tension will not be changed. So with the circulation, if the resistance in the arterial peripheral vessel is varied and the length of the pulsation depends on the time of fall in tension only, the pressure does not vary, if the vascular capacity is constant. THE FREQUENCY OF THE PULSE. 47 It is thus seen that the blood pressure need not depend on the arterial resistance, but if the pressure does not vary, the pulse-rate must do so. A desire to arrive at the genuine value of this theory of Marey’s, led me to make experiments similar to his own, as to the accuracy of his fundamental facts. My observations were divided into two series, to find,— 1st. Whether the pulse-rate was related to the capillary resistance. 2nd. Whether the pulse-rate depended on the pressure of the blood in the arteries. These points will be considered separately. Ist. The relation of the pulse-rate to the arterial resistance. The effect of exposing the surface of the body to the influence of different temperatures, whereby, as it has been my endeavour to prove elsewhere,* variations in the calibre of the eutaneous vessels are pro- duced, was carefully examined and the following tables embody my results, the curves being those of changes in pulse-rate. Ezperiment I. Temperature of the air 515° F. Nude at 11.57 P.M. Page 5. Lay down on floor, carpeted, on right side, at 11.58 p.m., with head on Nieut 50 55 60 65 ————_ foot-stool. Did not feel cold. Got up and put on night shirt and jumped into bed at 12.20; a skin glow came on at 12.29... Same position maintained in bed as when on carpet. Ezperiment IT. Temperature of air 50° F. Nude at 11.40 a.m. and lay down on right side. Experiment conducted exactly as the last. 11.45 12 NIGHT woe or i) ee bo Ww * “ Proceedings of the Royal Society,” 1869, p. 419. (Suprd, p. 6.) Page 6. Page 7. 48 THE LAW WHICH REGULATES Got up at 12 night, and in bed in Jess than a minute. A glow came on at 12.6. Experiment III. Temperature of air 52°5° F. To show that the change in pulse-rate did not depend on the effort of getting into bed. 11.10 P.M. 11.15 11.30 11.45 12 NIGHT Experiment conducted exactly as the first. Nude, lying on the right side, at 11.7 p.m. Got up at 11.26 p.m., put on night shirt, took it off again and lay down again on floor. Got up again at 11.45'5” and went into bed, in night shirt. A glow came on at 11.54'5” p.m. From these observations it is apparent that the effect of simply altering the condition of the cutaneous vessels, by varying their rela- tions to external agencies, varies the pulse-rate in a definite manner ; and thermometric results show that on warming the skin, as by covering it with bad conductors, the vessels are increased in calibre and the arterial resistance reduced. These experiments therefore show that reducing the resistance quickens the pulse. Marey’s own observations, specially as they are recorded mostly by the graphic method, are of themselves sufficiently convincing on this point. He compressed the abdominal aorta of a horse, per rectum, and found the pulse thereby rendered much slower. The same result followed compression of the human femoral arteries. The quickened pulse produced by the Turkish bath (in one case reaching the extreme rapidity of 172 in a minute on myself), is well known; as is the slow one following a cold bath, as shown by Drs: Bence Jones and Dickinson. From these many facts, all tending in one direction only, it may be stated that the rapidity of. the pulse varies inversely as the resistance to the flow of blood from the arteries. Qnd. The relation of the pulse-rate to the amount of blood in circula- tion, or to the blood pressure wm the artertes. The following experiments were made :— wet be 7 ~~ THE FREQUENCY OF THE PULSE. 4G Experiment IV. An old donkey which had been standing for more than half-an-hour in the room in which the experiment was conducted, had at 7.30 4.m.a pulse of 34a minute. At 7.40 half an ounce of chloral hydrate was given it in 2 oz. of water. At a minute 7.50 46 Standing unsteadily as if intoxicated. 7.52’ It fell down asleep. 755 43 7.59 40 : 8.8’ 48 A tap having been put in the jugular vein, but no bleeding having occurred. 8.12’ 52 Bleeding slowly from jugular. 8.15’ 67 3 64 Minute after minute, the animal having lost altogether a 62 about one pint or a little more of blood. » 60 8.19 59 Bleeding freely. 8.20’ 52 Bleeding ceased. 8.21’ 49 » ” 8.22’ 48 ” 2 8.22’ 30” Bleeding resumed. 8.23’ 30” 49 Bleeding. Resp. 11°5. 8.24 15” Bleeding ceased after loss of another pint and half. Page 8. 8.24’ 30” 43 Bleeding ceased. 8.25’ 15” 42 No bleeding. Resp. 11°5 8.26 42 a 8.28’ 15” 42 Bleeding freely. 8.30’ 30” 42 os » Resp. 13. 8.34 30” 37 3 “ 8.36 38 . ¥ 8.40 37 » ” 8.42’ 35 9 » Resp. 14. 8.45’ 36 % » Resp. 16. and from this time until 9.10, by which time more than half a pailful of blood had been lost and the carotid pulsations were very feeble, the pulse remained at 35°6 to 35 in a minute, with the respirations varying from 12 to 13 in the same time, and the loss of blood being continuous throughout. The animal did not move once through the whole experiment. Experiment V. A terrier dog had 30 grains of chloral given it in two doses, 15 grains first and another 15 grains about an hour after- wards. This did not render it quite comatose, so it sniffed chloroform until insensible, when a hemadynamometer was connected with one of _ its carotids, and a pressure of 6°6 inches was immediately registered, which was steadily maintained, undulating with the respiration. Page 9. Page 10. 50 THE LAW WHICH REGULATES Resp. Pulse in a minute. in a minute Pressure 40 186°5 6°45 inches In 8 minutes... 42 192 63 a » 2 x 40 188 64 » ” 2 ” ets 184 674 ” ae aoe 49 182 66, a 52 164 685, ee ee aba 159 7 x ” 1 ” ad 150 7 ” cS ee 70 138 a Sr ee = 142 67 oy, » 5 ” weed 149 68 ”» » 6 ” as 156 5'8 ” ” 7 ” =m 168 64 ” Ree agen aa 102 Sy ae » 10 » y= 129 21 ” ”» 5 ” bees 138 19 ” ee lg — 141 ae er ea ees 64 142 wt 3 S'< oa - 138 4 Seale In this experiment the bleeding occurred from the carotid, and took place between the pulse countings, which were traced on a re- volving drum. The great fall in pressure indicates the excessive bleeding in one case. In others a much less quantity of blood was lost on each occasion. There was no bleeding after the fall in pressure to 1°7 inches, and from that time the pressure and pulse became less and less till the animal died. Experiment VI. A rabbit was made comatose by 15 grains of chloral, and was bled to death, the whole operation taking half an hour. An hemadynamometer was connected with the carotid, and the blood was lost from the jugular of the same side, in drops con- tinuously. The pressure, at first at about 6 inches, fell at the end of the experiment, to less than 1 inch, when death occurred. The fol- lowing are the pulse-rates taken at equal intervals during the half hour :— In 10 seconds. In 10 seconds. 42°5 42°75 41°9 43° 44: : 42° 43° 42° 42°75 42°5 41° 415 A3° 42° Bleeding began.. 46° 42°4 44°5 41° 45°5 41°75 44:75 42° THE FREQUENCY OF THE PULSE. 51 In 10 seconds. In 10 seconds. 43°25 42° 44° 45° 43°5 43°9 43° 42- 43° SERShRE Experiment VII. A rabbit under the influence of 15 grains of chloral— When pressure 48 inches. Pulse 136 in a minute. ” 43 » » 133 » ” 28 » ” 127 ” ” 17 ” » 1382 ” ” 28 ” » 133 ” » 24 » 158 ” » 19 » » 144 ” ” rt » 133 ” ” “75 ” » 136 ” , ” 9 »” ”» 127 2? death from loss of blood. From these experiments it is evident that the pulse does not in- crease in frequency with loss of blood, as it did not do so in any one of them. ‘ In Experiment IV the pulse-rate rose on making the incision in Page 11. ____ the skin necessary to expose the jugular vein, and continued to do so q shortly after bleeding commenced, but soon diminished; and after ‘reaching 36 a minute, remained perfectly constant, notwithstanding a - continuous and considerable loss of blood from the vein until the. animal was almost exsanguinated. . zn With the rabbits the difficulty in keeping them completely under = the influence of the hypnotic, with the tendency to struggle, makes __ the results less uniform; but in all the cases there was a fall in pulse- rate, not a rise, accompanying the reduction in blood pressure. This fall, which was not very great, may result from the cooling of the _ surface, consequent on. the lessened circulation. __- From these observations it may be concluded that variations in the amount of blood in circulation do not vary the rapidity of the pulse, _ and consequently, that the pulse-rate is not dependent on the blood pressure as Marey supposed. The next question was—What law as to the frequency of the heart’s beats would satisfy these two aboye-proved facts, namely, the dependence of the pulse length on the arterial resistance and its non- dependence on arterial blood pressure ? ; The method adopted by Mr. Fleeming Jenkin for detecting the E2 Page 12, Page 12. 52 THE LAW WHICH REGULATES insulation of long cables at different times occurred to me as being subject to exactly similar laws, the time of fall of cable charge from tension to half tension, which he employs, varying directly as the leakage, and as that only. Can it be that the heart always recommences to beat when the tension falls a certain invariable proportion, and then only? This theory it was my next object to analyse, and the different elements into which it resolves itself were, and will be now, considered sepa- rately. Sao m First, as to the full meaning of the term—a uniform ‘insulation. A uniform circulation is one in which the quantity of fluid flowing through all cross sections of the circulating system is the same; for if the flow through one part were less than through another, there would be a tendency for the fluid to accumulate in front of the obstruc- tion, which is incompatible with the premises. As a consequence of this, the heart must always recommence to beat directly as much blood has left the capillaries as was sent out from it in the previous pulsation, and therefore the length of the pause or diastole must depend on the relative capacities of the heart and of the arterial system, and on the rapidity of the flow of blood through the capillaries. At this point the work of Poiseuille respecting the flow of fluids through capillary tubes is invaluable. He found* that, other things being the same, the flow of fluids through capillary tubes varies directly as the pressure. These results were verified by a Committee of the Academy of Sciences; and by an entirely different method I have been enabled to do the same on the vessels of the animal system. My method was the following in a particular case :—The kidneys of a deer, with the aorta and renal vessels intact, were removed from its body and placed for some time in water at 100° F.; the aorta was ligatured just below the origin of the renal arteries, and a uniform glass tube was tied into it just above them. Water at 100° F. was poured into the tube and it distended the organs; the tube was main- tained full by a continuous supply, which was suddenly stopped and the time of fall of the column from tension to half tension at different initial pressures observed, and it was always found that it took exactly the same time to fall from 40 inches to 20 inches as from 20 inches to 10 inches, thus verifying the law. This law being thus true, it is evident that if the capacity of the arterial system, including the left ventricle, varies directly as the * “ Recherches expérimentales sur le mouvement des liquides dans les tubes de trés-petits diamétres.”” ‘ Rapport de l’Académie des Sciences. Comptes Rendus.” Tome IV. 1842. THE FREQUENCY OF THE PULSE. 53 pressure, then the heart must always recommence to beat when the _ arterial tension has fallen a certain proportion; for with double pres- _ sure and consequently double amount of blood, the time of flow through the capillaries is constant, as the flow varies directly as the pressure; and with double resistance and unvaried pressure the time of flow is double also, for the heart pumps again when as much has gone from the capillaries as it has sent into the arteries and the rela- tive capacities of the heart and arteries do not vary according to the But does the capacity of the arterial system vary as the pressure ? This is a point which it is very difficult to prove. With regard to the heart, the following facts bear on it:—By connecting a syringe _ with the coronary arteries, or by tying it into the aorta and pumping Page 14. _ backwards, it can be shown that increasing the pressure in the coro- nary arteries increases the capacity of the ventricles. Also in many post-mortem examinations the heart is found with the ventricular cavities fully obliterated, and as they are not then in action, the capa- city of the heart and the pressure in it are at a minimum together. This is all the direct evidence that it is in my power to bring on this point. With regard to the arterial system and its capacity, the absence of blood in the arteries after death has been known from time imme- __—«moriail, and if their capacity varied directly with the pressure, it is ___ evident that that must be the case, both capacity and pressure being — ata minimum. A direct method of determining this point having occurred to me, the following description will illustrate it. In a rabbit one of the carotids was put in communication with a kymographion ; and during the time the recording drum was revolving, the chess was suddenly opened and the ventricles cut across transversely. The pressure fell rapidly to zero, and it is clear that the fall must have arisen from the escape of the blood through the peripheral vessels, as the aortic valves would close immediately. The curve of descent would take a definite form, which is easily expressed in mathematical language, if the capacity diminished as the pressure. ‘ Unfortunately the time required = to open the chest, and other difficulties connected with the operation, "prevented my results from being of much value, and Dr. Michael — Foster suggested to me that the same object would be attained if the Page 15. heart were made to stop by the action of the interrupted current on the pneumogastric nerve. Mr. Martin of Christ’s College, Cambridge, kindly sent me some traces thus taken, and one of the two which are suitable for measurement, entirely conforms with the law that the capacity of the vessels varies directly as the blood pressure, assuming Poiseuille’s law to be correct. The other curve does not exactly fulfil Page 16. 54 “THE LAW WHICH REGULATES the requirements, but varies very little from them. When further opportunity occurs, I hope to repeat these experiments on a larger scale. It can also be shown in other ways that the arteries do not obey the laws of ordinary elastic tubes. They are covered by a dense, scarcely elastic, fibrous coat which limits their distension, and they are sur- rounded by organs and muscles which are pressing on them in all directions. So it may be said at least that they do not vary in capa- city as simple elastic tubes, and that the difference is towards their varying directly as the pressure. However, the indirect evidence proves that the capacity of the arterial system, the ventricle included, varies directly as the pressure : for the facts above considered as to the frequency of the pulse depend- ing on the resistance, and not at all on the pressure, can only bé explained on this assumption. If the direct evidence ‘as to the dapat of the wiitiela had been contradictory, it is true that it would have been necessary to assume some error in the method of conducting the pulse experiments; but as above shown, it is quite in the right direction and only lacks partial direct verification. So much in the verification of theories connected with Physiology must depend on the way in which collateral facts are explained by ; them that it will be advisable now to consider some of them, and these considerations will be divided into two sections,—I1st. The explanation of the known variations in pulse-rate in health; and 2ndly, The expla- nation of the cardiograph laws. 1st. Variations in Pulse-rate in Health—With reference to these points, as on this theory change in pulse-rate can only depend on change in-arterial resistance, it is evident that Marey’s law will, upon his supposition as to the relation between blood pressure and arterial resistance, explain the phenomena equally well. The following are some of the best known :— The Effects of Respiration on the Pulse-rate——Physiologists, though not completely agreed as to the effects of respiration on the pressure of the blood in the arteries, all acknowledge that during inspiration the pulse quickens, and during expiration it gets slower, whether the pressure rises or falls. The theory under consideration clearly shows that this must be so, for during inspiration the expansion of the chest must reduce the pressure in the intra-thoracic aorta, and consequently its contained blood must fall in tension more rapidly than if the chest were motionless, and the more rapid tension fall causes increase in pulse-rate. In expiration the opposite occurs, diminution in chest capacity reduces the size of the aorta and consequently delays the time of fall of tension, and therefore slows the pulse. THE FREQUENCY OF THE PULSE. 55 If other remote effects of respiration tend to modify the pressure Page 17. in the vessels, it is evident that they would co-exist with the above and influence it but slightly, explaining the existence of the experi- mental discrepancies. The Effect of Position of the Body on the Pulse-rate.—The experi- ments of Dr. Guy led him to explain the differences in pulse-rate following change of position as depending on the amount of muscular effort necessary to maintain the positions assumed, and his explana- tion, assuming that muscular effort of itself can change pulse-rate, is very complete. It is curious that the theory under consideration gives an interpretation of the same facts, though very different from that of Dr. Guy. The following are the most essential facts :—The pulse is quickest while standing erect, slowest while lying, intermediate while sitting, slow while standing leaning, and while supported entirely, as by being bound to a wheel in any position. The following is the explanation :—While standing, the only soft parts of the body which support the weight of the body are the soles of the feet, and the weight is transmitted to them through non-vascu- lar and rigid tissues, cartilage and bone. Consequently the blood flows freely through almost all the vascular system unobstructed. But while lying, most of the weight is supported by highly vascular tissues, as the shoulders, arms, thighs, and legs, and consequently much of the circulatory system is greatly reduced in capacity from the compression ‘it experiences, and considerable resistance to the flow of blood is introduced into the system, the fall of tension is retarded, and the Page 18. pulse therefore rendered slower. In sitting, an intermediate condition is the result and an interme- diate rate of pulse is produced. Leaning while standing, and entire support on a wheel, both by introducing resistance from compression of soft parts, tend to make the pulse slow. Thus, according to Dr. Guy’s assumption, the slow is the normal pulse, and the quick the induced; upon the fall of tension theory the reverse is the case. The occurrence of bed-sores and the paleness of a compressed part prove that pressure disturbs the uniformity of the circulation. The rapid pulse after a meal, during digestion, depends on the relaxation’ of the vessels of the alimentary canal while its functions are being performed. : 7 4’ aie! —S Ad . Jk ile el a aT nL = fa e 4 In a paper on cardiograph tracings* from the human chest wall, * Since writing the paper referred to, a further comparison of tracings has shown me that in the slow pulses taken while lying, I mistook the primary systolic Page 19. Page 20. 56 THE LAW WHICH REGULATES published in the 5th vol.of the “ Journal of Anatomy and Physiology,’’* I have endeavoured to substantiate a law respecting the elements of the heart’s beat, which may be thus enunciated :— The heart’s beat consists of two parts, which for any given pulse- rate do not vary in their ratio to one another; but the length of the first part varies inversely as the square root of the sap of the pulse. A second series of measurements of the cardio-arterial intervals, published in the ‘‘ Proceedings of the Royal Society,”’t have further verified the law just stated, and in the rest of this paper it will be assumed as proved. No theory respecting the circulation throws light on its significance; but the one which it has been my endeavour to demonstrate above gives a very satisfactory explanation of it, which will now be considered in detail. First, the heart’s beat consists of two parts, which for any given pulse-rate do not vary in their ratio to one another. It having been proved previously that the pulse-rate does not depend on the blood pressure, and, as shown now, the length of the first part of the heart’s beat not varying when the pulse length is constant, it is evident that the length of the first part of the pulse-beat does not depend on the - blood pressure in any way. - Again, the first part of the pulse-beat is compound, for it is thee interval between the commencement of the cardiac or ventricular systole and the closure of the semilunar valves; therefore it may be divided into the systole and the valve-closure interval. Physiologists have laid very little stress on this valve-closure interval, it generally being considered as instantaneous. But in the study of cardiograph tracings it is to be remembered that the dis- tances between events occurring within one-fiftieth of a second of one another can be appreciated withont much difficulty, and there is every a priort reason for believing that this interval has a longer duration than that. In my paper on the cardiograph trace, reasons have been given for the belief that in quick pulses the commencement and the end of this valve-closure interval are indicated by separate and distinct changes of direction in the curve, and its length as obtained by measuring from these points agrees entirely with that required from arguments to be mentioned further on. It may be called the diaspasis, rise for the auricular, and was so led to the conclusion that the length of the cardiac intervals depended in some measure on the position of the body. This is incorrect, as subsequent measurements show me, and the length of the first part does not vary with the position of the body ; the proper equation for finding the cardiac first part under all circumstances being zy = 20 V2. * Supra, p. 18. + Supra, p. 32. THE FREQUENCY OF THE PULSE. 57 that is, the period during which the heart is being opened out by the regurgitation of blood from the arteries. The length of the combined systole and diaspasis not depending at all on the pressure, and it being constant for any pulse-rate, it is infinitely probable that the systole and diaspasis separately are inde- pendent of the pressure, and this is extremely interesting, as it gives a further insight into the mechanism of the heart. For, in order that the duration of the diaspasis should not vary with different blood pressures, it is evident that with higher pressures there must be greater obstruction to the heartward flow of blood, otherwise the valyes would then close more quickly. And this is exactly what would be expected from the combination of Mr. Bryan’s observations concerning the shape of the heart, and Briicke’s theory of the active diastole of the ventricles.* According to the latter author the cardiac Page 21. "muscular tissue has no inherent power of opening out the ventricles, but remains inactive after systole, during diaspasis in fact, until the regurgitation from the aorta has closed the aortic valves and so un- covered the orifices of the coronary arteries, immediately upon which the resulting sudden turgescence of the heart’s walls makes them open up. Mr. Bryan has shown that during systole the whole heart alters its position as a result of its change in shape during contraction, and recovers it during diastole; therefore the greater the force of contrac- tion the more will it alter its shape, and the more difficult will it be for it to resume the original one, which has to be done partly by the regurgitating arterial blood; but the greater the blood pressure, the greater will be the facility for overcoming this greater work, which two, as must be the case, vary together. This argument explains how the diaspasis need not vary in length with different blood pressures. Next, with regard to the systole. As the first part of the heart’s beat varies as the square root of the length of the beat, and as the diaspasis, a part of that first part, does not vary with the blood pres- sure, upon which alone it can depend, it is necessary that the other component of that interval must vary more than as the square root of the pulse length. And to find how much more quickly, it is necessary to obtain the actual length of diaspasis. Careful measurements of a cardiograph trace, beating 102 in a minute, give the ratio of the systole to the whole beat as 1 to 3°1915, and that of the first part to Page 22. the whole beat as 1 to 20, which leaves the ratio of the diaspasis to the beat as -187 to 1, or the diaspasis length as 00183 of a minute. A similar length of diaspasis is found from quicker pulses. % * Mr. Bryan’s paper is in the “‘ Lancet,” Feb. 8th, 1834. - Briicke’s theory appeared in “Sitzungsberichte der Wiener Akad. der Wiss.,” 1854, Vol. XIV. p. 345. A paper on the same subject by myself will be found in _ the “ Journal of Anatomy and Physiology,” May, 1869. (Supra, p. 3.) Page 23. 58 THE LAW WHICH REGULATES The great interest attaching to these figures is that, when with the diaspasis equal to ‘00183 of a minute, the ratio of the systole to the diastole is enquired into, it is found that there is a very simple rela- tion between them, and that after subtracting this diaspasis of con- stant length, there remains the systolic varying as the square root of the diastolic period, and with no other diastolic length is so simple a ratio obtainable, which is all-important, because it will be seen that the systole must depend directly on the previous diastole. Next, considering the systole itself; the fact above demonstrated, that its length does not depend on the blood pressure, is extremely important, and can only be explained by assuming that when the pressure rises, the circulation through the coronary vessels increases to a sufficient extent to enable the heart to get through the extra work it has to perform without altering the duration of its action, or, in more precise terms, the nutrition of the walls of the heart must vary directly as the blood pressure in the aorta. But the systolic length varies as the square root of the diastolic, in other words, the longer the time of nutrition of the heart, the longer the systole. This at first sight seems an anomaly, but the theory that the pulse-rate depends on the fall in tension only, presents a most complete explanation, and so throws great light on cardiac action in general. Consider the heart as a pump working against a certain pressure, and filling an elastic reservoir with a certain resistance to the outflow of its contents. Varying the pressure has been shown to have no effect on the lengths of the different parts of the pulsation for the reasons given above; and it has next to be considered how it is that varying the resistance changes the lengths of the elements of the revolution. This pump, directly its muscular fibres begin to contract, exerts its full pressure, for there is nothing to prevent it doing so. But during the previous diastole it was supplied by blood at a certain definite pressure and for a definite time, both of which factors limit the force of the systole. Consequently, the ventricles produce directly their full systolic pressure, and maintain that pressure until they are empty. But it is evident that the time necessary for emptying them of a definite amount of blood under these conditions must depend on the rapidity of the flow from the capillaries, for when the flow is halved the systolic time must be doubled, if no other force come into play ; in other words the length of cardiac systole is a function of the arterial resistance; and the pulse-rate has also been shown to be a function of the same, upon the fall of tension theory. It has been proved that the systole varies as the square root of the diastole, not directly with it, as might be supposed. This clearly shows that the time of diastole influences the length of the systole THE FREQUENCY OF THE PULSE. 59 and shortens it, in other words, strengthens the heart, according to Page 24. the law that may be stated thus: the nutrition of the heart varies as the square root of the time during which the coronary circulation is maintained. It will strike some as peculiar that no mention has yet been made of the influence of the nervous system on the heart. But it appears to me that the facts which have been brought forward have not called for any special reference to it. May not the law it has been my endeavour to prove, be but an expression of that action in the healthy body ? For it must depend on a somewhat complicated mechanism, as is shown by the fact that it is almost impossible to contrive a self-acting engine which would pulsate in accordance with its require- ments. As is well known, the effect on the kymographion trace of slightly stimulating the pneumogastric nerves is greatly to amplify the oscil- lations, and at the same time to lower the mean pressure; while cutting the pneumogastrics produces the reverse effects. The larger oscillations of the hemadynamometer column in the former case show that the proportionate tension fall and the time of pulsation are both greatly increased, and from previous considerations it is evident that these are necessarily associated when, as now, no influence is being exerted on the peripheral vessels.* Further, these amplified oscillations must be attended with an Page 25. abnormal enlargement of the ventricular cavities during diastole, for the time intervening between the beats being increased, the amount of blood which flows through all segments of the circulation between any two pulsations must be also more considerable. Having arrived so far, it is extremely interesting to observe how an augmentation in the degree of cardiac dilatation during diastole, as a cause, will include and correlate all the peculiarities which are observed when the depres- sor nerve is thus operated on; and it is not unreasonable, therefore, to suppose that this is the direct effect of its action. As the quantity of blood contained in the heart at the end of diastole has been shown to depend on the circulation through the coronary vessels, it is evi- dent that the explanation of any variations in the capacity of the ventricles must be referred to changes in the cardiac walls themselves. Just as the degree of rigidity of an india-rubber tube through which a current of water is flowing, can be made to vary by changing the diameter of the orifice from which the fluid is allowed to escape, so the turgescence of the ventricular walls, or what is the same thing, * In rabbits the normal fall of tension as judged by the hemadynamometer trace is about =5th of the whole, while when the pneumogastric is stimulated it may increase to th or more. Page 26. Page 27. 60 THE LAW WHICH REGULATES the amount of active diastole of the heart, can be altered, by varying the diameter of the small arteries of the coronary system, their con- traction producing a greater, and their dilatation by facilitating the flow of blood through the capillaries a less, degree of diastolic enlarge- ment of the ventricular cavities. . From the above argument, therefore, the amplified range of pres- sure and time depending on change in heart capacity, and the change in capacity being caused by modification in the calibre of the smaller coronary arteries, it is almost a logical necessity that the function of the pneumogastric nerve is to regulate the degree of tonicity of those vessels, and Dr. Brown Séquard, from entirely different facts, has also published it as his belief that the pneumogastrics contain fibres which contract the small coronary vessels.* It will be noticed that throughout this paper it has been assumed that the systole never recommences until the ventricular cavities are completely filled, that is, until a pressure equilibrium has been arrived at in the interior of the heart. Perhaps it is the absence of pressure which admits of the heart recontracting, but this is a doubtful point, and until more is known as to the mechanism of muscular action in general, it is probable that the question as to the reason why the heart recommences to: beat ata particular moment will remain unsettled. Sir J. Paget,f when he pointed out the relation of rhythmic nutrition to rhythmic action of nerves and muscles, laid the foundation for a scientific treatment of the subject, and the law which it has been my endeavour to substantiate, is only a precise method of expressing that relation. The following summary of the main features in the circulation, as they appear to me, may assist in explaining some of the previous arguments. The circulation of the blood is maintained by the repeated con- traction of the heart. Hach cardiac revolution is divided into three parts, the systole, the diaspasis, and the diastole. The following laws hold with regard to the length of these intervals. I. The systole together with the diaspasis, or in other words, the first cardiac interval, varies as the square root of the whole revolution. IJ. The systole varies as the square root of the diastole. III. The diaspasis varies very slightly with different pulse-rates. The amount of work that the heart has to perform in maintaining the circulation depends on two sets of changes which may occur in * See “ Principles of Human Physiology.” By Dr. Carpenter, 1869, p. 219. Foot-note. + “Croonian Lecture.” Royal Society, 1857. ee 6: - the system; 1. “Variations i in the blood pressure. 2. Variations in the resistance to the outflow of that fiuid from the arteries. As the capacity of the arteries, including the ventricles, varies directly as the blood pressure, and as the flow of blood from the capil- - laries does the same, the frequency of the heart’s beats is dependent on the resistance to the capillary outflow, and not at all on the blood pressure ; in other words, the heart always recommences to beat when the blood pressure in the x eras arteries has fallen a certain inva- riable proportion. Variations in blood pressure result from: 1. Absorption into, and excretion from, the vascular system of fluids. 2. Changes in the capacity of the arterial system, which occur on the contraction or relaxation of the muscular arteries. 3. Changes in the amount of Page 28. available blood, which result from the hemastatic dilatation of some of the yielding vessels on altering the position of the body. As changes in the first of these cannot be very sudden, and those in the latter are never very considerable, the mean blood pressure in health varies but little during short intervals. Variations in peripheral resistance result from: 1. Different de- grees of tonicity or patency of the muscular arteries. 2. Different resistances in the venous system. The former may occur indepen- dently in one or other system of vessels, as the cutaneous or the alimentary ; also mechanically from pressure on a part of the body. The latter are insignificant in health. The heart depends for its power of doing work on chemical pro- perties in the blood it pumps into the systemic vessels, and as the blood reaches it direct from those vessels, the cardiac intramural circulation varies with the changes in the former; and the length of the systole varying only as the square root of the time of diastole, the degree of cardiac nutrition varies directly as the systemic blood pres- sure, and as the square root of the diastolic time. The coronary arteries supplying the whole heart, the work done by the right ventricle is governed by that done in the left; thus the supply of blood in the left auricle is always rendered sufficient for the require- ments of the systemic circulation; though, as there is no reason for believing that the resistance in the pulmonary vessels varies with that of the systemic, there must be some peculiarities in the former circu- Page 29. lation (which may explain the variations in the ratio of the number of pulse-beats to respirations, in some cases). The auricular contraction is a very small force, and its function is most probably to close the tricuspid and mitral valves. The heart commencing its systole as a whole, it is highly probable that the impulse for action is given by a force which affects both 62 LAW REGULATING THE FREQUENCY OF THE PULSE. ventricles ; such is found in the coronary circulation and the active diastole produced by means of it. In conclusion I have to present my best thanks to Dr. Michael Foster, Professor Sanderson, and Professor Pritchard of the Royal Veterinary College, for the opportunities they have afforded me in trying the experiments above detailed; without their assistance it would have been impossible for me to have put the law of the relation of blood pressure to pulse frequency on any satisfactory basis. June 10th, 1872. ON SPHYGMOGRAPHY. 63 10. ON SPHYGMOGRAPHY.* Page 399. Part L Ir is to the happy conception of Marey’s, in which he obtained correct amplification of sudden small movements by employing the elastic force of a spring rather than the statical pressure of weights, that we owe the form of Sphymography, now well known to most students of ; physiology ; and the introduction of this fresh method of research has given a stimulus to the progress of precise thought respecting the cir- culation of the blood and the action ©f the heart, which must neces- sarily prove of extreme value in the investigation of pathological conditions, now quite beyond our grasp. Marey’s original instrument, as constructed by Breguet, is so well known, and so many excellent descriptions are accessible,t that it is not necessary to go into detail regarding it. In most respects it is the best form that has been introduced, and much credit is due to M. Breguet for the excellence of the mechanical construction. This maker has lately introduced a second instrument, which has the advan- tage of removing the chief defects of the original one, and of intro- ducing very little intrinsic error. The chief objection to the old, or knife-edge construction, as it may be called, from the fact that the recording lever is connected with the pulse pad by means of a sharp steel edge, depends on the necessary sliding of that knife-edge on the steel surface below the recording lever, by which means the distance between the axis of rotation of the lever and the point of contact of the knife-edge must vary with every movement of the former; the accompanying diagram shows how this is the case for two different heights of the lever. In consequence of this imperfection, two traces taken, one near the top, and the other lower down on the recording paper, present cha- Page 400. racteristics which are in many respects different, and whose differences arise solely from the sliding above mentioned. The lever in the upper trace falls less rapidly than in the lower, as the knife-edge has to slide over a much larger surface in the former than in the latter, where it scarcely moves at all. Asa result of this, the length of the first part of the higher trace is apparently greater than it ought to be. * Part I, “ Journal of Anatomy and Physiology,” VI. pp. 399-404. May, 1872. Part IT, lc. VII. pp. 98-105. Nov. 1872. + Marey, “Physiologie médicale de la Circulation du Sang,” Paris, 1863; B. F. Foster, M.D., ‘‘ On the use of the Sphygmograph in the Investigation of Disease,” 1866; J. Burdon Sanderson, M.D., F.R.S., “ Handbook of the Sphygmograph,” 1867 ; and others. Page 401. 64 ON SPHYGMOGRAPHY. * Fig. 1. TSEREREI TAVERRAA TRE TIRAHATT IOS Another objection is, that there is no firm connection between the pulse-spring and the recording lever, so that when the latter is sud- denly raised, as by the systolic impulse, the contacts of the lever, inter- mediate brasswork, and pulse-spring, are not maintained, and a “ per- cussion-rise”’ is seen in the trace. This error is partly obviated by a small spring, which presses on the recording lever, and tends to keep it in contact with the other moving parts, though this is omitted by some, much to the detriment of the instrument. Though in the taking of the trace itself, this absence of connec- tion between the lever and pulse-spring is disadvantageous, yet in the applying and removing of the instrument it is of extreme value, for it allows of any amount of pressure being put on the latter, without pro- ducing any strain or injurious effect on the former. In practice this quality is invaluable, and it is almost impossible to obtain it in any construction other than the one under consideration. Breguet’s new sphygmograph is very simple and ingenious in prin- ciple, and in practice works admirably. As to the older instrument, the recording lever is fixed to the body of the apparatus by an axis or arbor, but the novelty consists in the manner in which this is brought into connection with the pulse-spring. The lever is of the third kind, as in the other instrument, but it has no steel surface below it for a knife-edge to play on, and it is, when not in use, entirely disconnected from the pulse-spring. Fig. 2. Near the middle of the arbor there is a small circular ring of brass, ON SPHYGMOGRAPHY. 65 surrounding it, which has a diameter of about one-sixth of an inch. On its outer surface this ring is grooved and an endless screw is cut in this groove. : There is no brasswork attached to the pulse-spring corresponding to that in the older instrument. In its stead a long flat slip of brass, fixed about two-thirds the length of that spring from its attached end, extends onwards over the ivory pad. Just above this pad a steel screw about an inch long, with a milled-head at one end, is fixed at the other to the unattached extremity of the piece of brass just mentioned, by a hinge, and is so arranged that by means of the pulse-spring it is retained either at right angles to it, or lying along it, on the same principle that an ordinary knife-blade can be fixed half open or closed. This long screw, when at right angles or nearly so, to the spring, comes in contact with the endless screw cut in the brass ring surround- ing the arbor, and as they are of corresponding size, they bite and are retained in contact by the pulse-spring pressing on the fixed end of the screw, which is squared off in such a manner that it shall continue to press slightly backwards when in contact with the ring. It is evident from this construction that any up or down movement communicated to the pulse-pad, produces a corresponding rise or fall in the recording lever. When not in action the long serew can be thrown out of gear, Page 402. down on the spring, as seen in Fig. 2, and it has also a movement -which allows of its being turned round, which can be employed as a fine adjustment in regulating the height of the lever point. Tn this instrument the distance between the axis of rotation of the arbor and the part of the long serew which is in contact with the ring surrounding it, cannot vary, whatever the distance of the pulse-pad, and therefore no error is introdueed in that direction. Also the diffi- culty in disconnecting the spring from the axis, after the trace has been taken, is very slight; though it requires some little practice to do this, before removing the instrument from the wrist, as it always should be. This, the rackwork sphygmograph, will probably supersede the kmnife-edge one by degrees, for the tracings are more uniform, and in other respects quite as good as those obtained by the earlier instrument. For cardiograms it is not so advantageous. With regard to the best means of binding the sphygmograph on the arm, the original method adopted by Marey, of lacing it with a silk ribbon to the side-lappets, is as efficient as any. The wrist should be always bent a little backwards, and care should be taken that the pad presses on the lower end of the radial artery, not on the super- ficialis vole, as is apt to occur if itis fixed too far forwards. The pad introduced by Mr. Berkeley Hill enables the correct position of the hand to be maintained with facility, but it is scarcely necessary, as a F Page 403. 66 ON SPHYGMOGRAPHY. little practice removes all difficulty. It is worthy of notice that better traces can be obtained after the instrument has been applied a few minutes than immediately; for the pad, by its pressure on the skin, drives blood from the small vessels covering the artery, and so lessens the distance between it and the pulse-spring. To record the movements of the lever, it is best to employ highly glazed paper which has been smoked thinly by the flame of a compo- site candle; and to effect this properly, the paper must be folded on the metal plate, which is connected with the running gear of the sphygmograph, and moved quickly up and down in the flame. By this means its edges are not charred, and a more uniform film is pro- duced. The pen must have a fine sharp point. A spirit varnish, such as is used for photographic plates, fixes the tracings, if a little is poured over them gently, and the paper is subsequently warmed. An entirely novel form of sphygmograph has been constructed by M. Longuet,* which, from a figure in Dr. Loraine’s work, appears to possess all the requirements of an accurate instrument, except in the recording portion, where the pen, instead of writing laterally, as it might easily be made to do, rests on the paper horizontally, and has to change its relation to the body of the instrument whenever the pulse- pad moves in the least. In principle its construction is in a great measure similar to that of the cardiograph in my combined cardio- sphygmograph, which was described in this Journal in May, 1871.+ It has a dynamometer attached, which measures the pressure and its changes throughout the beat. One of the points in which Marey improved upon Vierordt’s original sphygmograph was, that he made his lever write laterally instead of at its tip, by which means he obviated all difficulties connected with an exact up and down movement of the recording pen. But this gives rise to another slight error, which can and must be corrected in com- paring the length of the elements of the pulse-beat. The lever, when the clockwork is stationary, traces a curved line and not a straight one, the curve being part.of a circle, of which the lever is the radius, and its arbor the centre. Therefore, when the watchwork is in action, the horizontal relation of points at different heights on the recording paper is not truly represented by straight lines, drawn perpendicular to the length of the paper, but by lines projected from these points in curves parallel to that produced by the lever on the paper when it is stationary. These lines can be easily formed after the tracing has been removed from the apparatus, by scratching on it with a needle that is tied to a nail, or a piece of board with two pieces of cotton of the * “ Bulletin de l’Acad. de Médecine,” XXXTIT. p. 962, 1868. + Supra, p. 27. I i taal dbs ON SPHYGMOGRAPHY. 67 length of the recording lever, one to its eye, and the other near its point, if the tracing be fixed with its lower edge in the same straight line as the nail; for then the paper and nail bear the same relation to Page 404. one another that the former did to the arbor of the lever when it was being recorded. Dr. Sanderson,* Mr. Mahomed,+ and others have introduced differ- ent ingenious methods for regulating and measuring the pressure applied on the artery, which they have described in full in their books and papers on the subject; but the desired results can be only approxi- mate, as from variations in the elasticity of the skin in different individuals and in the same individual under different atmospheric conditions, complications are introduced which cannot be eliminated. The sphygmograph is a bad hemadynamometer at its best, and as such its employment will probably diminish. A glance at the work of Dr. Lorainet will show that numberless variations in the character of a tracing can be introduced by very small changes in the position, &c., of the subject experimented on. But there is one part of the trace which is not affected in any way by these complications, and that is the relation borne by the length of the different portions of each beat to one another. Whether the arm is raised or lowered, whether the pressure is great or small, the interval between the systolic rise and the commencement of the diastolic rise does not vary in the least. These intervals cannot be correctly estimated, except by the aid of the sphygmograph; and as the value of any method of investigation is greatest in that direction in which it is least influenced by surround- ing circumstances, it is, as it will be my endeavour to prove when considering the tracings themselves, for the measurement of these intervals that we must look for the future value of the sphygmograph. Parr II. Ty endeavouring to form a correct estimate of the significance of the Page 98. various details of the sphygmograph trace, it will be necessary to enter somewhat minutely into the consideration of each of the several mutually related forces which, by their combined action, produce the resulting curve. As some of these forces are but little understood, it is clear that any attempt to explain the pulse movements by argu- _ ments deduced from tracings obtained from a “ schema” of the circu- lation, can only be of value, as far as they relate to forces acting on A £5 * Loc. cit. + ‘ Medical Times and Gazette,” 1872. t “ Etudes de Médecine Clinique. Le Pouls.” Paris, 1870. . F 2 Page 99, 68 ON SPHYGMOGRAPHY. the “schema” which are strictly comparable with similar forces at work in the human body. This consideration has prevented my basing any deductions on facts derived otherwise than from the animal body itself. To commence with, it will be necessary to describe the tracings obtained by means of the “ Hemadromograph” of Chauveau. Dr. Lortet, of Lyons, has given an excellent figure and description of this instrument,* which, from its simplicity and efficiency, is a perfect masterpiece of mechanical design. By means of two levers it indicates on a revolving drum both the modification in the diameter of an artery during a pulse-beat, and also the changes in the velocity of the blood-current at the same spot, simultaneously. The former of these results is obtained by the use of an ordinary sphygmoscope, an instru- ment which measures indirectly the modifications in the area of an artery ; and the latter by a long lever which, at its attached end, after passing through a slit in the side of the tube, projects a short dis- tance into the vessel, which allows of its being moved backwards and forwards by the blood-current. The accompanying trace (Fig. 1), taken with this instrument from the carotid of the horse, is copied from Dr. Marey’s work.t Fig. 1. Hemadromograph trace from the carotid of the horse. a, The current trace, in which the horizontal dotted line is the zero, indicating no current in either direction. B. The sphygmoscope trace. In both, the vertical line 1 cuts the curve at the commencement of systole, the line 3 at the moment of closure of the aortic valve, and the line 4 when the centri- fugal current recommences. The figure reads from left to-right. * ** Recherches sur la Vitesse du Cours du Sang dans les Artéres du Cheval.” Par M. L. Lortet, M.D. ‘ Annales des Sciences Nat.,” Tome VII. Zoologie, 1867. An earlier but very similar instrument, by M. Chauveau, is described in Marey’s “ Physiologie Médicale de la Circulation du Sang,” 1868, p. 156. + Loc. cit., p. 273. ON SPHYGMOGRAPHY. 69 The upper curve («) is that of the blood-current; in it the horizontal ‘dotted line indicates the zero, or line of no current in either direction; all above shows a centrifugal current, and all below a heartward stream. The velocity of the blood’s movement is measured by the distance of the point which is under consideration from the zero line in the trace. The lower curve is that obtained by the sphygmoscope, and is in all respects strictly comparable with an ordinary sphygmo- graph tracing. The slightly curved vertical lines which intersect both tracings are drawn by the levers when the watchwork is at rest, and the points where similarly situated lines intersect the pulse traces are exactly simultaneous, to indicate which they are drawn. With regard to these figures, Marey makes the following observations :— “I. The commencement of the pulsation coincides with the produc- tion of a rapid centrifugal current. II. The summit of the pulsation is not reached before the centrifugal current has already ceased. Ill. At the instant of closure of the sigmoid valves, a retrograde current is flowing in the carotid, as indicated by the position of the curve, which at that moment is below zero. IV. After the closure of the sigmoid valves a fresh centrifugal current originates, which consti- tutes the secondary pulsation—(dicrotism).” Dr. Lortet explains the portion of the upper or current-trace which is below the zero line, by Page 100. _ supposing that the shock of closure of the sigmoid valves is sufficiently “great to produce a stretching of the proximal end of the aorta and of ; the valves themselves, which results in a retrograde current for a short time in the large arteries after the valves have closed. Marey’s expla- nation, as far as it goes, seems much more satisfactory, and it will be necessary to discuss it more fully. Before doing so the precise defini- tion of some of the terms employed will not be out of place. The main rise in the sphygmograph trace may be termed the primary rise, and that which occurs just after the small rise and fall which is nearly simultaneous with the sinking of the current-trace below the zero, and which therefore commences with the secondary centrifugal current, may be termed the secondary rise; it is almost always very clearly indicated in sphygmograph tracings (c, a, Fig. 2). The interval be- tween these two rises may be called the sphygmosystole, for it is the ie time during which the systole at the heart influences the pulse-beat; Page 101. it corresponds to the first part of the pulse-beat in my former com- munications, and it must be remembered that it is not synchronous with the cardiac systole, as 1 have endeavoured to show elsewhere.* From a study of tracings obtained by the use of the cardio-sphyg- mograph I have demonstrated, in the paper just referred to, that in 2 7 * “ Proceedings of the Royal Society,” 1871, p. 318, e¢ seg. (Supra, p. 32), and “ Journal of Anatomy and Physiology,” May,1871. (Supra, p. 27.) 70 ON SPHYGMOGRAPHY. Fig. 2. Sphygmograph tracings, all taken from the same individual in health, under different conditions, to show the effect of difference in pulse-rate on the trace. a. Pulse-rate 44 in a minute. B. ” 63 ” 33 Y: ” 72 ” ” 8. y 206, é. ” 137 ” ” mN = ” 172 ” ” They are all drawn on one scale and read from left to right. the sphygmograph trace of slow pulses the closure of the aortic valve occurs when the lever is at the lowest point of the notch that is nearly always present in the sphygmosystole, which is also clearly seen in the lower of the two tracings in Fig. 1, the curved line 3 cutting it exactly at the place which represents the moment when the valve shuts. Consequently, in the upper trace, the line 3 must also cut it at the same time, and this occurs when the retrograde current is at its maximum, as would be expected, the valve being closed by the regurgitating blood. But it may be asked, how is it that the back- ward current between the lines 3 and 4 in a, Fig. 1, is associated with a rise in the sphygmoscope trace? and to explain this clearly it will be necessary to refer to some of the elementary principles which operate in the transmission of currents through elastic tubes. First, it can be demonstrated that increase in the diameter of an artery may originate from two quite independent sets of causes, one being the simple result of the heart sending more blood into it in a given time than it can dispose of, the other being a shock-expansion, comparable with the waves of condensation and rarefaction in the air, which con- stitute sound. Most of the important elements of the pulse-trace are ON SPHYGMOGRAPHY. 71 referable to the former of these causes, as is proved by the general simi- larity between the two curves « and 8, in Fig. 1. But when there are rises in the second, which are quite independent of changes in the first, they must evidently be the result of some sudden movement in the circulating system, originating a shock, which is not of a character to affect the current in any way. It is but reasonable to suppose that such should occur as a result of the closure of the semilunar valves, and the rise and fall between the vertical lines 3 and 4 in the lower Page 102. trace (8) of Fig. 1, commencing immediately on their closure, and being unconnected with any current changes, can be nothing else than a shock-wave. _ At first sight it would seem probable that when the semilunar valves had closed, the retrograde current in the aorta would immedi- ately cease; but that such is not the case is clearly proved by the continuance below the zero line of the current curve for a short period after that event (between the lines 3 and 4in the upper trace). It is by the combined operation of two causes in the same direction that » this result is produced. First, the orifices of the coronary arteries being quite close to the semilunar valves, it is evidently necessary that the blood which enters them during diastole must be derived from the aorta, and so tends to produce in it a retrograde current. Secondly, the equilibrium of the arterial system is disturbed during the closure of the aortic valve, for immediately the systole has termi- nated, the only force tending to prevent the blood from regurgitating into the heart is the statical resistance of the ventricular walls, which at that moment are closely approximated, causing their cavities to be completely obliterated. This resistance is clearly much less consider- able than that offered by the heart-walls during the systole, one being a statical and the other a dynamical condition; consequently the arterial blood rushes back, pushing asunder the ventricular walls, and in so doing developing a sufficiently rapid retrograde current to close the semilunar valves. The interval thus arising, namely, the time between the end of systole and the complete closure of the aortic valve, I have called the diaspasis, in a pamphlet on “On the Law which Regulates the Frequency of the Pulse,”* and there is consider- able evidence to show that when the pulse-rate does not vary it is con- stant; also that it varies very slightly with different frequencies of heart-action, occupying in slow pulses about 0°002 and in quick ones 00018 of a minute.t The diaspasis is so short in duration that it has Page 103. 2 ; : f * Published by H. K. Lewis, 139, Gower Street, London. (Supra, p. 45.) t+ In my pamphlet on “ The Law which regulates the Frequency of the Pulse,” from a mistake on my own part, a statement is made as to the length of the diaspa- sis, which is incorrect. Mathematical friends soon informed me of my error, as, Page 104, 72 ON SPHYGMOGRAPHY. terminated before the loss of blood from the proximal aorta into the heart is made good by any retardation of the onward current, or a reverse stream in the more distant vessels originates; consequently this restoration of equilibrium has to be in great measure effected after the valves have closed, encroaching on the diastole to form the second cardio-arterial interval of the cardio-sphygmograph trace, which does not cease until all the complications attending the closure of the semi- lunar valves have come to an end, whereupon the again augmenting centrifugal current, or in very low-tensioned pulses, such as that figured in Fig. 1, the diminishing centripetal one, originates the secondary rise in the sphygmoscope trace. In both the primary and secondary rises of the pulse-beat (Fig. 1), it is found on inspection that the summits of the pulsations (8) are delayed upon, or are not reached so soon as the centrifugal current maxima (@); and that such should be the case is essential, in a cireu- lation maintained by an intermittent motor organ, which, like the when precisely stated, the problem is a. manifest contradiction. In p. 22 the diaspa- sis is discussed, and in the summary of the results arrived at, in p. 27, the following are given as the relations of the lengths of the various elements of the pulse-beat to one another. 1. The systole together with the diaspasis, or in other words, the first cardiac interval varies as the square root of the whole revolution. 2. The systole varies as the square root of the diastole. 3. The diaspasis is constant. The incompatibility of these three statements it is not difficult to prove, for if the first and second are true, the third cannot be so, and, as a fact, the diaspasis is not constant. There is every reason to believe that for a given pulse-rate the diaspasis does not vary ; and, on subtracting its length, as obtained by measurement in quick cardiograph tracings, from the first part of slow ones, I found that the remainder, the systolic interval, varied very nearly as the square root of the diastolic. On repeating the operation on the pulse of intermediate rapidity a similar result was obtained, and the error being extremely small, I attributed it to my not having extracted the necessary square roots to a sufficient number of decimal places, and thus felt justified in making the generalization given above. Since the mistake has been pointed out to me I have repeated the arithmetical computations more care- fully, and find that what I had first supposed were errors on my part, are constant variations, which prove that, the other statement standing as above, the diaspasis is slightly longer in slow pulses, occupying approximately 0°002 of a minute in a pulse of 61, and 0°0018 of a minute in a pulse of 152 in a minute. This fact, therefore, leads to the conclusion that the rapidity of the fall of blood-tension has an influence on the length of the diaspasis, lengthening it slightly when the tension-fall is retarded, probably because the previous systole is then more powerful and gradual. It is to be noted that the second cardio-arterial interval of cardio-sphygmograph tracings is almost of the same length as the diaspasis, and varies in the same or in a very similar manner, which may be the cause of the somewhat undecided nature of the notch in the sphygmosystole of slow pulses. From these remarks it is necessary to substitute for statement 3, as given above, the following: 3. The diaspasis varies, being slightly longer in slow pulses. ON SPHYGMOGRAPHY. 73 heart, rests between its pulsations; for after the rush of blood into the arteries immediately the sigmoid valves open, during the rest of the systole the blood which leaves the heart is employed in retaining the higher pressure in the vessels, as will be explained more fully further on. We are now ina position to consider the human sphygmograph tracing from the wrist, and on looking at Fig. 2 («), which is from a slow pulse, it is evident that in all respects it closely resembles that taken with the sphygmoscope from the carotid of the horse, which has been discussed above, and there is every reason to believe that the details originate from similar causes. The primary rise (a) is followed by a gentle fall; this is soon broken by the shock-wave (4) consequent on the closure of the aortic valve, and is followed by the secondary rise (c), which commences when the centrifugal current is augmented by the recommencing onward current in the aorta. The sphygmo- diastole is remarkably uniform and uninterrupted. Between the tracings of slow and quick pulsings there is at first sight not much resemblance, but it is not difficult to obtain a series between them exhibiting every intermediate condition (Fig. 2, a, 8, 7). The most important cause of the variations exhibited by pulses of dif- _ ferent rapidities, is that the ratio between the length of the sphygmo- ~ systole and the sphygmodiastole is not constant. For instance, when the pulse is 114 a minute, the sphygmosystole occupies just one half the beat; but when it is 40 a minute, it only occupies one quarter of the whole revolution. It is evident that this must influence the general appearance of the trace, and as the length of the sphygmo- systole never varies in health for any given pulse-rate,* a knowledge of the ratio of the length of sphygmosystole to that of the whole beat is sufficient datum for determining the pulse-rate. In the paper just referred to I have given several measurements of these ratios, and have shown that the length of the sphygmosystole maintains a very definite relation to the length of the beat, varying as its cube root, consequently when the length of the revolution increases in the series 1, 8, 27, 64, Page 105. &c., the sphygmosystole only does so as 1, 2, 3, 4, &c., so that if we call the rapidity of the pulse z, and the number of times that the a sphygmosystole is contained in the beat y’, the length of the sphyg- mosystole can be found from the equation ay’ = 47 3/2. Further, the cardio-sphygmographf shows that the interval between the closure of the aortic valve and the commencement of the secondary * “Proceedings of the Royal Society,” No. 120,1870. (Supra, p. 14.) + “Journal of Anatomy and Physiology,” May, 1871. (Supra, p. 27.) Mae 74 ON SPHYGMOGRAPHY. radial rise (the second cardio-arterial interval) varies but little with different pulse-rates, while that between the primary radial rise and the closure of the aortic valve (the conjugate cardio-arterial interval) does so much more rapidly, both being longer in slow pulses. This also greatly influences the appearance of the pulse-trace, for, as pre- viously shown, the small rise and fall at the end of the sphygmo- systole results from the shock of closure of the aortic valve, and as this occurs in slow pulses an appreciable time after the primary rise has reached its maximum, it is clearly seen as a separate element of the curve. But in quick pulses the second cardio-arterial interval is nearly as long as in the slow ones, while the conjugate cardio-arterial interval is much shorter, consequently the shock-rise and fall follow- ing the aortic valve closure is thrown back, as it may be termed, on the primary rise, and, being blended with it, is not separately distin- guishable. This is the cause of the simplicity in the sphygmosystole of quick pulses. THE SOURCE OF NERVE FORCE. 75 11. ON THE SOURCE OF NERVE FORCE: A THEORY.* Page 251. THE universally acknowledged inefficiency of any theory at present before the scientific world to account for the origin of the force by which, through the intervention of the nervous system, organs at a distance from one another are placed in communication, makes me feel justified in publishing a theory which has been in my mind for some time, and which during that time accumulated information has not in any way shaken. ' The following is my proposition:—The species of energy which exhibits itself in the form of “ nerve force,” is electricity of thermo- electric origin, resulting from the fact that the surface of the living animal body is always colder than its interior. In other words, the available energy resulting from the interior of the living animal body being at a higher temperature than that of the surrounding medium, is expended wholly, or in great measure, in generating the force called — nerve force. In the discussion of this problem, the following are the considerations on which it is based. 1. The temperature of the interior of the living body is always "greater than that of its surface, because all animal life is only a form of chemical degradation, and is therefore necessarily attended with evolution of heat; which, no other force intervening, will always keep the interior of the body hotter than the medium in which it lives. When the temperature of the air in higher animals exceeds or nearly approaches that of the body, which varies but little, special arrange- ments (perspiratory, &c.) come into play to diminish that of the surface. 2. There is an available source of energy in the body, which has been but little considered by physiologists, depending on the tempera- ture of the surface being lower than that of the interior. The theory propounded gives employment for this force. r 3. In the struggle for life, the individuals that economise -the forces at their disposal are most likely to survive; it is therefore im- Page 252. probable that this not inconsiderable source of energy should have escaped employment in this struggle. 4. The actual construction of the nervous system is quite sufficient for the working of such a force as the one proposed. In addition to the already known properties of the nerves, including their good con- ducting power, it is only required that a thermo-electric current be cap- * “ Journal of Anatomy and Physiology,” VII. pp. 251-4. June, 1873. Page 253. 76 THE SOURCE OF NERVE FOROE. able of being generated between soft tissues of different composition or structure, such as the extremities of the sentient nerves and the corium in which they are embedded. Although Magnus was not able to produce any thermo-electric current between fluid metals, his experiments are not in any way con- clusive against the possible existence of such currents between dif- ferent tissues ; the subject is in a position for actual demonstration, no doubt, and I much regret that as yet I have not had opportunities of attempting to prove or disprove it. Notwithstanding this, some of the circumstantial or collateral evidence is so strong, that, without any direct proofs, I feel justified in assuming its correctness. The follow- ing points strongly favour my theory. 1. Within certain limits, which are those ra which the body is most generally exposed, the energy of the individual (which must be closely related to the supply of nerve force) is greater as the tempera- ture of the air is less. On a cold day, in a cold air, there is more will and power to work physically than in hot weather, in a hot air, during which languor is a prominent feature. In a paper published in this Journal* I have proved that the circulation through the skin in man varies according to the temperature of the atmosphere, and that when 70° F. of the air is reached, perspiration commences. Consequently by this means a difference of about 30° F. is always maintained in health between the surface and the interior; and exposure to the highest temperatures of the Turkish bath, when continued for some time, does not disturb this condition. But a hot-water bath is fol- lowed by very different consequences; if the body and face be im- mersed in one of 100° F., and breathing be performed through a tube projecting from the mouth to the surface, as I have tried, faintness, or loss of nerve power, comes on very rapidly, and is it not because the temperature of the surface is raised to that of the interior? At all events this explanation is as reasonable as any other. This faintness is soon recovered from on the reapplication of cold to the skin. 2. The effect of muscular exercise is to raise the temperature of the body, and so to increase the difference of temperature between it and the external air; and when great muscular exertion has to be sus- tained, as in rowing, most of the clothes have to be removed in order to allow of the rapid cooling of the surface, the necessity for which is keenly felt. 3. During intra-uterine life the active chemical changes going on in the body of the foetus maintains its temperature at a slightly higher point than that of the mother; but immediately after birth the nerve force generated by the cooling of the surface brings the intercostal * Supra, p. 87. THE SOURCE OF NERVE FORCE. 77 and other muscles into play, and the child is otherwise much more active than when in utero. 4, The source of the body heat being central, and the circulation of the blood tending to render that of all the parts uniform, it is evident, as a simple physical deduction, that the temperature of the portion of the body which corresponds to the heated end of the thermo-electric couple (which in the theory now under discussion can only be the brain), must be lower than that of the blood, because the heat supplied to, and developed in it (the brain), is partly employed in generating the electric current which is circulating. Dr. John Davy noticed that the temperature of the brain of the rabbit was pecu- liarly low, considerably below that of the abdominal viscera; his re- sults have been considered improbable by some, but have never been refuted ; they are strongly in favour of my theory, which, as shown above, explains them completely. Upon my theory the mechanism of the nervous system may be thus summarised— The afferent nerves are the conductors to the nerve centres of the electric current which is generated by the contact of their peripheral ends with the tissues of the cooled skin, which they supply. The brain is the largest of the centres towards which the nerve current is . directed, the other ganglia forming the smaller. Through these centres the currents, as through an elaborate commutator, are split up or concentrated’ in a manner not understood as yet, to be directed Page 254. along the efferent nerves, which are always so situated as to be beyond the reach of external cooling influences. Where an organ acts in any way automatically, it. generally has centres of its own, of a size vary- ing in degree according to its automaticity, and these minor centres are only to a certain extent subject to the influence of the brain. As in the working of the electric telegraph, no return or second special conductor is required to carry back the current to the point from which it started; for where an efferent nerve terminates in a muscle, it loses its insulatiug covering, and so is put into indirect com- munication with the peripheral sentient nerves through the inter- vention of the mass of body tissue generally, which, though its resistance is much greater, offers an merpeay larger mass to be traversed by the current. Page 140. Page 141. 78 ON POINTS CONNECTED WITH THE 12, ON SOME POINTS CONNECTED WITH THE CIR- CULATION OF THE BLOOD, ARRIVED AT FROM A STUDY OF THE SPHYGMOGRAPH-TRACE*+ — (Pl. III.) Sivcz my first communication to the Royal Society, “On the rela- tive Duration of the Component Parts of the Radial Sphygmograph- Trace in Health” (“Proceedings of the Royal Society,” XVIII. p. 351+), it has not been my good fortune to find any similar observa- tions by other physiologists, either in favour of or in opposition to my statements. From that time my attention has been continually directed to similar phenomena; and the employment of similar methods has led to results which seem to have an important bearing on the problem of the action of the heart. It is evident that a thorough knowledge of the nature of the pulse in the arteries, when combined with that acquaintance with the anatomical mechanism of the heart and arteries that can be arrived at from post mortem examina- tion, is sufficient basis for a fairly thorough study of the circulation of the blood. It has been my endeavour, by the employment of the sphygmograph as constructed by M. Marey, to obtain an amount of information from the curves which it produces sufficient to generalise on the nature of the cardiac action in some of its details which have not as yet attracted attention. The results will be stated in the form of propositions. Prop. I. The length of the interval between the commencement of the ventricular systole at the heart and the closure of the aortic valve does not vary when the pulse-rate is constant, and varies as the square root of the length of the pulse-beat—being found from the equation ay = 20 /x, where « = the pulse-rate, and y = the ratio borne by the above-named part to the whole beat. This law, in a somewhat modified form, was enunciated by myself in 4 paper published in the “Journal of Anatomy and Physiology” (V. p. 17),t where the peculiarities of the curves taken in the lying posture misled me as to the point of commencement of the ventricular systole, and led me to state that posture had an effect on the duration of the systole. Such, however, is not the case; for, while lying, the weight of the heart is apparently sufficiently great to neutralise the * “ Proceedings of the Royal Society,” XXIII. 1875, pp. 140-51, Pl. 5. Read April 23, 1874. An abstract of this paper is published, loc. cit., XXII. pp. 291-3. + Supra, p. 14. t Supra, p. 18. CIRCULATION OF THE BLOOD. 79 effect on the trace of the auricular contraction, and to make the thus taken trace deficient in the rise which at other times results from that contraction. Atall events if this assumption be made, itis found that the lengths of the different parts of the beat are not influenced by posture, and they agree exactly with the above-stated law. The following are measurements made since the publication of the original paper, which tend fully to confirm the above statement :— vce | Tames | ict, the first ° Ww. is eeactend ta the beat on formula whole beat. ay = 24/2: 46 2°925 2°93 43 2-8, 2°88 2-885 49 2°85 2-86 52°5 2°71 2-765 56 2°63 2°675 57 2-75 2-66 58 2°65 2-625 60 2°63 2°59 64°5 2-556 2-49 69 2°45 2-4 74 2°28 2-325 79 2°23, 2-275 2°24 80 2 -24375, 2-207 2°225 81°5 2-2, 2°185, 2-093 2-2 St 2-105 2°175 85 2°09 2°16 86 2-17, 2-053 2-155 88°5 2-245, 2-275 2-11 905 2-062 2-1 92 2°12 2°09 < 92 2 -0875 2-08 94 2°14125 2°05 Prop. Il. The length of the interval between the commencement Page 142. of the primary and the dicrotic rises in the radial artery is constant for any given pulse-rate, and varies as the cube root of the length of the pulse-beat—being found from the equation zy’ = 47 3/z, where 2 = the pulse-rate and y’ = the ratio borne by the above-named part to the whole beat. This law was enunciated in the paper before referred to as read before this Society by myself, and published in its ‘‘ Proceedings” (XVIII. p. 351).* Since that paper was read a fresh series of measurements have strongly confirmed its accuracy, and practice in manipulation has diminished my limits of experimental error so far that a difference of 5 per cent. from the calculated results is rarely found. * Supra, p. 14. Page 143. 80 ON POINTS CONNECTED WITH THE The following table contains some of the more recent results and one or two more careful measurements of old traces :— Ratio borne by sphygmosystole to whole beat, as Pulse-rate. F Calculated from ‘ound : — by measurement. equation xy’ =47 x (approximately). 38 4175 4°18 43 °5 8 °825 3°8 44°5 3°7875 3°75 56 3°29 - 3°22 58 3°195 8-185 59 3 °185 3°0 63 2°911 2°96 63°5 2 °938 2-95 64 2 °904 2°93 65 2°83, 2°821 2°9 67 2°825, 2°788 2°84 66°5 2 °889 2°795 69 2°7 2°78 "3 2°625 2-685 105 2°132 2°13 140 1-785 1°75 Prop. Ill. The length of the interval between the primary and the dicrotic rises follows the same law in the carotid and posterior tibial that it does in the radial artery. That such is the case as far as the femoral and posterior tibial arteries are concerned is shown by Dr. Galabin in a paper “On the Secondary Waves in the Pulse,” recently published (Journal of Anat. and Phys., 2nd series, No. xiii). It is not necessary, in proving this law, to undertake any large series of measurements; for if all those which are taken agree exactly with the calculated result obtained from the radial equation, the probability that it is correct is almost infinitely great. In a boy, etat. 16, whose radial pulse was previously proved to follow the above law exactly, the following are the results obtained by measuring the carotid tracings :— Ratio borne by sphygmosystole to whole beat, as Pulse-rate. Found Calculated by measurement. from radial equation. 67 2-899 2°84 68 2 ‘827, 2°6 2°8 72 2°7144, 2°7 77 2 +583 2°59 77 2 °594 2°59 CIRCULATION OF THE BLOOD. sl In another subject, stat. 22, the following are the results :— Ratio borne by sphygmosystole to whole beat, as Pulse-rate, Found Caleulated by measurement. from radial equation. 77 2 6625 2°595 78 2°575 2-575 85 2-443 2°44 . With regard to the posterior tibial artery, most of the results were obtained by the employment of the double sphygmograph, to be de- scribed further on, im which the superposition of the simultaneous posterior tibial trace on that from the radial artery showed that the interval between the commencing primary and dicrotic rises is the same in both. The following are a few independent measurements from tracings from the artery behind the ankle :— Proportion borne by first os — by Pulse-rate. part to whole hole ay a beat in ankle trace. bf in radial trace (approximately). 70 2-7 2-76 73 2°675 2-685 80 2-596 2-525 82 2 °4575 2-5 82°5 2°517 2-495 88 2°378 2-378 Corollary.—The length of the interval between the primary and secondary rises being exactly the same in the carotid, radial, and pos- terior tibial arteries, which are three vessels at very different distances from the heart, it is evident that the length of this interval is constant throughout the larger arteries, and must be of the same duration at the origin of the aorta that it is in the radial artery at the wrist. The corollary to Prop. III leads to theoretical results of consider- Page 144. : able importance ; for as the duration of the different elements of each 3 beat in the radial artery is the same as that in the commencing aorta, a by superposing the sphygmograph-trace upon the cardiograph-trace at . any given pulse-rate, a comparison can be made between the duration of the different physiological changes going on in the heart and those going on in the commencing aorta; in other words, the time during which the ventricular and arterial systoles are continuous can be ascer- tained with precision by an indirect method, which alone is possible in the human subject. G Page 145. 8 ON POINTS CONNECTED WITH THE Taking the equations given in Prop. I and Prop. IT, the length of the systolic portion of each beat in the cardiograph- and sphygmo- graph-traces may be calculated with facility for any value of . From the equation above given, namely, zy = 20 /x and ay’ = 47 (/2, it is found that the length of the arterial systole is shorter than the car- diac, as would be expected, because the cardiograph-trace is an indica- tion of the movements in the muscular walls of the heart, and not of the contained blood, and because a certain tension must be reached by the intraventricular blood at the commencement of the systole before it can push open the aortic valves. The sphygmosystole being therefore shorter than the cardiac systole, it becomes a question, when an attempt is made to superpose them exactly, as to whether they correspond at the commencement or the end of the cardiosystole. This is easily answered; for independent obser- vations show the points in both at which the semilunar valves of the aorta close. These points in the traces must evidently be simultaneous, which is therefore the same thing as saying that the interval between the greater cardiosystole and the shorter sphygmosystole is at the commencement of the cardiosystole. This interval, the existence of which is well indicated in Marey’s cardio-aortic tracings from the horse, may be termed the syspasis (the time during which the ven- tricles are raising the pressure of their contained blood); and the following Table, obtained from the two equations just mentioned, gives its length at different pulse-rates :— ‘0018753’ at # = 36 approximately. 00132986’ ,, « = 49 Ks 000931’ ,, w = 64 ns 00004199" ;, « = 81 Xe 0003766’ ,, « = 100 ia 00024645’ ,, « = 121 ie 000118’ —s,, x= 144 000000’ Ss, # = 170 > From this Table it is evident that the syspasis varies considerably with different rapidities of pulse, decreasing rapidly with increase in the pulse-rate and becoming nil when it is 170 a minute, which may be fairly conceived to be very near the limit of cardiac rapidity in man. That this interval (the syspasis) should vary so considerably in length with different pulse-rates is not easy to explain at first sight; nevertheless a careful review of the different processes which are in operation in the heart at the time has suggested to me an explanation which seems reasonable. It depends on the fact that the extreme shortness of the diastole makes any variation in its length have a CIRCULATION OF THE BLOOD. 83 marked influence on the amount of blood which enters the capillaries of the walls of the heart, and consequently infiuences the amount of work which the muscular fibres of the ventricle have to perform in emptying their interstitial vessels before they can commence con- tracting on the blood in their contained cavity. Experiment shows that the rapidity of the pulse does not depend on the pressure of the blood in the arterial system ;* consequently the length of the syspasis is not influenced by the arterial blood-pressure, which is the same thing as saying that the force of the cardiac contraction varies directly as the blood-pressure; for then the muscular power of the ventricular walls to overcome the intramural distension, varying with it, prevents its duration from being modified. It has been my endeavour to show elsewheret that the force of the heart’s contraction is modified by the length of diastole, varying as its square root. Such being the case, it is evident that the length of the syspasis must vary with that of the diastole, though not tothe extent that is found to occur. But the diastolic period being always so short, it is evident that the longer it is the more thoroughly does the heart-tissue get permeated with blood, in a way which can have little or no influence on its nutritive power, but a great effect in modifying the length of the syspasis in the direction which is found to occur. Again, referring to the results of the cardio-sphygmograph obser- vations published by me in the “ Proceedings” of this Society (vol. xix, p. 318), that paper contains a table of the length of the different cardio-arterial intervals; and if from the first cardio-arterial interval, as there defined, the length of the syspasis be subtracted at the cor- responding rates, it will be found that the remainder of the interval is of exactly the same length as the second cardio-arterial interval, which, on the assumptions made, it could only be, as both the systole and the shock of the closure of the aortic valve are propagated along the arteries from the same point under similar circumstances. The following table gives the lengths of the first cardio-arterial interval from which that of the syspasis as above determined has been sub- tracted, and by their side the lengths of the second cardio-arterial interval, as copied from the table in the communication referred to; their similarity cannot be the result of simple coincidence, as they are derived from independent sets of measurements. * “Journal of Anatomy and Physiology,” November, 1873. (Supra, p. 51.) + Ibid. vol. viii. (Supra, p. 59.) I (Supra, p. 32). Page 146. 84 ON POINTS CONNECTED WITH THE First cardio-arterial Second Pulse-rate. interval eardio-arterial with syspasis subtracted. interval. 36 ‘00238982’ ‘00239821’ 49 -00233314’ 00233342’ 64 002274 00227425’ 81 *00220541’ -00220546’ 100 *0021875’ *00218745/ 121 *002084.55/ *0020847’ 144 -0020185’ ‘0020185’ 170 *0019704/ *0019729’ After the completion of the cardio-sphygmograph tracing above referred to, it was my endeavour to obtain satisfactory double sphyg- mograph tracings from arteries at different distances from the heart. Two or three unsuccessful attempts suggested the plan which has proved successful. It was soon evident that there is only one artery, other than the radial, which it is possible to manipulate with any degree of facility, especially when the experimenter is the subject of experiment. This artery is the posterior tibial at the ankle, where it runs in the interval between the internal malleolus and the tuberosity of the os calcis, just before it gives off the internal calcaneal branches. On myself, this artery is as superficial and as easily reached as the radial; in the sitting posture it is quite under command when the foot is crossed over the opposite knee; it is considerably further from the heart than the radial; and to obtain as great a difference as pos- sible, the right wrist was on all occasions the one experimented on, the wrist and the ankle being, as far as can be estimated on the living body, 29 inches and 523 inches respectively from the aortic valves. Before going further it will be necessary to consider the sphygmo- graph-trace from the posterior tibial artery at the ankle. Wolff* has published the results of his observations on the dorsalis pedis artery ; and as they correspond with those from the ankle-trace of the pos- terior tibial, they may be recapitulated. He remarks that the pulse at the foot has a general resemblance to that at the wrist, it differing in the primary ascent being less abrupt and the summit less acute. In the descent the secondary undulation is remarkably insignificant. The other minor undulations are less constant. My observations con- firm the above with respect to the general similarity between the two pulses, the greater obliquity of the primary rise, and the less constant character of the minor undulations; the secondary rise has, however, * “ Characteristik des Arterienpuls.” CIRCULATION OF THE BLOOD. ~ 85 | never struck me as peculiarly insignificant, though it has peculiarities, to be mentioned. immediately. The ankle-trace of a pulse at about 70 a minute, as taken with an ordinary sphygmograph, differs from that at the wrist in more than Page 147. one point. The primary rise, as previously mentioned, is less abrupt ; the following fall is more considerable, and is not broken by the notch nearly constantly seen in wrist-traces of this rapidity. The secondary rise starts from a lower level and is well marked, reaching its climax considerably nearer the next primary rise than in the wrist-trace. There is, however, another feature in the early part of the secondary rise in ‘the ankle-trace, which deserves special attention because of its general occurrence. As is well known, in wrist-traces the secondary rise com- mences promptly and is quite uniform in character, but in ankle-traces there is nearly always a short horizontal continuation of the curve im- mediately following the primary fall, the point of departure of the two lines being clearly indicated by an abrapt, though not consider- able, change in direction. This horizontal portion of the trace is not of any considerable length, being im a pulse of 70 a minute about one- eighth of the whole beat; it is followed by a well-defined secondary rise, which is much longer and more gradual than the primary. Though described above as horizontal, this short interval between the two undulations is not so always, being frequently slightly oblique, sometimes in one direction, sometimes in the other. When its curve is downwards (that is, when it tends in the same direction as the primary fall), it may appear to be part of that event, which would then look as if broken; when its curve is upwards (that is, when it tends in the same direction as the secondary rise), it makes the trace appear more normal in comparison with that from the wrist. Having now explained the ankle sphygmograph-trace, in consider- ing the simultaneous wrist and ankle traces, it will be necessary to commence with the description of the instrument employed to obtain them.- A drawing from it above is seen in Plate III, fig. 1, from the side in fig. 2, and a double sphygmogram is given in fig. 4. The double sphygmograph is constructed from two of the ordinary sphygmographs of Marey, as first constructed by Breguet. One, that ‘employed in taking the ankle-trace, retains all its original parts, except the side lappets for fixing it to the arm, and its recording- apparatus receives the double trace. A second lever is fixed in con- nexion with it by two uprights so placed as to allow the axis of the second lever to be parallel to and above the one belonging to the instrument, sufficient room being left to allow the latter to move un- obstructed up to the top of the recording paper. This second lever, which is a fac simile of that used in the sphygmograph, is placed so Page 148, 86 ON POINTS CONNECTED WITH THE that it will write on the same recording-paper as the first; but its position is reversed. The accompanying sketch (fig. 3) will show this point, it representing a side view of the ordinary knife-edge lever upside down—that is, with the surface (s) on which the knife- edge ought to slide uppermost. the object of this arrangement will be seen immediately. The second sphygmograph has the jes wieeke removed, as well as the brasswork which is fastened to the spring that presses on the pulse, to the end of which a small wire loop is soldered. In addition, a small piece of wood is screwed into the nearer of the two holes by which the watchwork was fixed, in such a way that it can be made to revolve with difficulty. The two instruments are fastened together by means of a screw and nut in the foot-sphygmograph, which bind a brass plate in that for the wrist. This screw is fixed on a plate of brass which is attached to the end of the instrument furthest from the watchwork in the manner shown in the figure. The brass plate in the other sphygmograph, which it binds, is fixed on the side of the body of the instrument close to the arbor of the lever. The exact position of these additional pieces of brasswork has to be determined by the direction that a silk cord takes when, fixed at one end to the arbor- end of the inverted lever mentioned above, it is threaded through the loop on the tip of the spring of the wrist-sphygmograph. This cord has to be parallel to the sides of the ankle-sphygmograph, when the two instruments are fastened together with the nut at right angles to one another. On commencing to take a double trace the nut is unscrewed, and the two instruments are separated from one another. The wrist- sphygmograph is then bound, as usual, on the rightarm. The silk cord attached to the arbor-end of the wrist-pulse lever (the upper one in the ankle-sphygmograph) is then threaded through the loop at the tip of the wrist-spring, and the binding-screw to fix the two instru- ments is passed into the hole in the plate of the wrist-sphygmograph made to receive it; after which, the nut being screwed fast down, the two sphygmographs form a single mass. The silk cord is then carried round the piece of wood at the watchwork end of the wrist-sphygmo- graph, and, after being slightly tightened, is fixed in a groove on its side. The whole is now ready for commencing the trace. To do this the ankle instrument (with that for the wrist attached to it and to the arm) is placed over the left foot, which has to rest on the right knee, parallel to the direction of the leg, with the watchwork towards the body. The recording-paper is placed in position; the silk thread is tightened by slightly turning the wooden peg to which it is fixed, and the wrist-lever is made to pulsate by it towards the upper part of the recording-paper. The ankle-sphygmograph, held by its watchwork CIRCULATION OF THE BLOOD. 87 end in the left hand, and attached at the other extremity to the right wrist, is then pressed down on the inside of the left foot (which rests on the right knee), in such a way that its pulse-pad compresses the posterior tibial artery where its pulsation is most manifest. The lever is made to record on the lower part of the smoked paper, below the one connected with the wrist. When both levers are found to be working freely, the recording-paper is set’ moving by liberating the watchwork-catch with the left thumb, which is close to it. The respi- ration must be checked during the time the recording-paper is moving, to prevent irregularities in the trace. Results arrived at from the Study of the Simultaneous Wrist and Ankle Page 149. Tracings. In employing the tracings obtained from the above compound in- strument, two objects were kept in view—irst, to find the interval between the commencement of the primary rises in the wrist and ankle curves; and secondly, to observe whether or no the superposition of the one trace upon the other verified or falsified the statement made in Prop. III, that the lengths of the different parts of each element of the curve were the same in the two arteries. 5 The following table contains the measurements of the lengths of the intervals between the commencement of the primary rise in the wrist and ankle tracings at different rapidities of pulse, from which it is clear that this interval varies very slightly within the range that can be obtained, and that the tendency is for it to be very slightly longer in the slower pulses. Rapidity of Length of interval between commencement of systolic rise pulse. at the wrist and at the ankle. 62 00115’ occurring 14°08 times in each beat 63 00125’ »? 12°7 ” 2 2 67 001343’ z rst ee Sanat ¢ > 0013278’ 2 11°24 ” ” 2» 70 001222’ ” 117 2? ” 23 71 00136’ n 102 » ” ” ” 00124 »” 11°41 2? 5 2 . 0013’ 108 =, nies 72 0012’ ” u7 »” ” » 2 -001206' ss 1152 =, Gees 79 001145’ 2? 11-06 » ” 2» < so 00126’ »” 9 96 ” ” 2 81 001233’ »” 10 37 ” >»? 7 82 001123’ Se 10°86, as ~ 33 00122 » 10 3” »” 3 95 00122’ ” 8 67 ” bP 2 98 “001085' » . aE » » 99 00116’ 3? 8 607 »” 2? 2? Page 150. 88 ON POINTS CONNECTED WITH THE which gives an average length of ‘0012314 of a minute for all the rates. It being possible to estimate with considerable accuracy the dis- tance from the aortic valves of the spots on the arteries at which the instrument is usually applied, it becomes a point of interest to deter- - mine from the facts arrived at the rapidity with which the primary undulation travels from “its origin (the heart) to the peripheral vessels. The radial artery at the wrist and the posterior tibial artery at the ankle are, as nearly as can be determined, 29 inches and 52} inches respectively from the origin of the aorta in myself (on whom all the tracings have been taken), as previously mentioned; and as the time of transit of the wave varies very little with different rapid- ities of pulse, a single example may be taken to illustrate the point in question. With the heart beating 100 times in a minute, the time taken by the primary wave in reaching the wrist (that is, the length of the first cardio-radial interval with the syspasis subtracted) has been shown in a previous table to be ‘0021875 of a minute. Adding to this the interval between the radial and ankle primary rise at the same rapidity, which is very nearly ‘00116 of a minute, ‘0033475 of a minute is the time taken by the systolic wave in travelling from the heart to the ankle. But if this wave went the extra distance to the ankle (52°5 —29 =) 23°5 inches, at the same rate at which it reaches the wrist, the length of the first cardio-malleolar interval would be ‘00459375 of a minute (29 : 52°5: : 21875 : 459375); but it is only ‘0033475 of a minute, which is considerably less; consequently the wave augments in rapidity as its gets further from the heart, a phenome- non beyond my power to explain. By superimposing the wrist-trace from a simultaneous sphygmo- gram on that from the ankle, it is found that the components of each are of exactly similar duration, though the peculiar short interval following the dicrotic notch in the latter sometimes complicates the results. This exact similarity in length of the different elements of the two pulses is not, as will be found by those who attempt to measure them practically, self-evident from the tracings themselves ; because the one being slightly later than the other, and the watch- work varying in rapidity, gradually increasing and then declining, the radial, which is the earlier, is slightly the shorter in the com- mencement of the trace and the longer towards its end. In the middle of the recording-paper the two coincide. It may therefore be said that the compound sphygmograph-trace is entirely in favour of the correctness of Prop. IIT. In conclusion, the following is a summary of the results arrived at in this communication :-— I. The lengths of the different elements of the pulse-beat being CIRCULATION OF THE BLOOD. the number n being theck of 90 CIRCULATION OF THE BLOOD. the same in arteries at different distances from the heart, the radial sphygmograph-trace expresses their duration in the aorta. : II. The cardiosystole being longer than the sphygmosystole at all possible pulse-rates, the excess in the length of the former expresses the time required by the heart to reach, from a state of rest, a systolic pressure sufficient to open the semilunar valves. This interval, termed the syspasis, is constant for any given rapidity of cardiac action, and rapidly decreases as the pulse gets quicker, becoming nil at a rate of 170 a minute. III. The interval between the commencement of the primary pulse- rise in the radial and that in the posterior tibial artery is less than would be estimated from the time taken by the same wave in travelling from the aortic valve to the radial artery. The woodcut (p. 89) will assist in illustrating the mutual rela- tions of the different component parts of the cardiac revolution, as its different elements are there shown in their actual relations one to the other. $ THE TELSON OF THE MACRUROUS CRUSTACEA. 93 13. ON THE TELSON OF THE MACRUROUS CRUSTACEA.* Tue relations of the telson of the lobster and its allies are so variously regarded by zoologists of the present day, that no apology is needed in bringing forward any facts which tend to settle the point. By Milne-Edwards it is considered as a seventh abdominal segment ; but I cannot find in his writings any reasons given for his belief. Van Beneden is also stated by Professor Rolleston to hold the same opinion. Professor Huxley considers the telson an azygos appendage, and not a true segment of the body; and Professor Rolleston agrees with him, stating that it only carries appendages in one or two cases, whereas it is a law common to all Crustacea, that every segment has its ap- pendages. An attempt will be here made to show that in a specimen of Scyllarus arctus, in the Zoological Museum of the University of Cam- bridge, there is sufficient evidence to prove that the telson is a true body segment, and that it is provided with true segmental append- ages, though the nature of these is somewhat modified by cohesion and adhesion. In this specimen the sizth abdominal segment is in the main similar to that in the lobster, but its dorsal surface is grooved instead of plain. The infero-lateral terminations of its dorsal shield are slightly recurved and not sharply pointed, those of the first abdominal segment being decidedly so, but the acumination becomes less marked in each succeeding one. The swimmerets are greatly developed, the propodite not exhibiting any decided spinous protuberances. Both the exo- and the endo-podite, which are expanded horizontally, are composed of two, a proximal calcified and a distal fin-like portion ; the anterior margin of the former in each of these segments being prolonged outwards in the form of a spine. The distal fins are com- posed of a translucent membranous substance supported on a radiating framework. At the attached extremities of the anterior and posterior margins of these fins, there are small elongated calcified masses, which seem to be the points at which their delicate structures come in contact with one another and the neighbouring parts. Ventrally there is a transverse calcified bar, concave forward, * “ Journal of Anatomy and Physiology,” V. pp. 270-3. May, 1871. Page 271. Page 272. Page 273. 94 THE TELSON OF THE MACRUROUS CRUSTACEA. slender, and composed of a central and two lateral elements, which are fixed to the sides of the dorsal shield. The seventh segment, or telson, consists of a distal fin and a proxi- mal calcified portion. The fin is azygos, semi-elliptical, and supported on rays, like the exo- and endo-podites of the sixth segment. It also resembles them in having two lateral elongated calcified masses at its margins. There is no median separation. The calcified portion consists dorsally of a small semicircular cen- trum or dorsal shield, which is close under the dorsal plate of the preceding segment, and is separated from the lateral masses by a marked groove, not by an articulation. These lateral masses are each distinctly separable into three parts: 1st, a thin longitudinal one, which approaches the sixth dorsal plate, and ends posteriorly in a sharp spine. Internally it comes in contact with the next, the 2nd part, which latter joins the azygos centrum at its antero-internal margin, and at its distal end gives attachment to the 3rd part. Beyond the centrum there is a considerable interval between the two lateral masses dorsally. The 3rd part consists only of a free spine, directed backwards and articulated to the 2nd. It rests on the azygos fin. : Ventrally this segment presents only a short transverse bar, in front of the anus, composed, as in the preceding one, of a central and two lateral portions. On the supposition that this seventh segment is a true one, the ~ small dorsal centrum corresponds with the much larger one in the other segments, while the swimmerets are represented by the lateral masses described above; the 2nd part of which corresponds with the propodite, and the lst and 3rd with the exo- and endo-podite respec- tively, each bearing spines, and connected with the propodite and with the fin. The short transverse ventral bar corresponds to the narrow cen- trum, and is composed of the portions, as in the preceding segment. I have no means of telling whether it is laterally connected with the dorsal shield. The position of the anus behind this seventh transverse bar is strongly in favour of the alimentary canal being segmentally terminal, although this condition is disguised by the coalescence of the lateral appendages above, which consequently makes the anus ventral. If the above explanation is accepted with regard to this particular species, it is highly probable that it is true in those allied to it, and would then refer to the lobster and cray-fish. Professor Rolleston says of the latter,* that the proximal segment of the telson is not * “ Forms of Animal Life,” 1870, p. 95. THE TELSON OF THE MACRUROUS CRUSTACEA. 95 calcified continuously across its ventral surface, as the other segments are; but this appearance would arise from the small size of its real centrum, which, as in Scyllarus, would be quite anterior, and the © coalescence in the middle line of the proximal portions of the endo- podites of the segment. In the telson of the lobster, by looking at its dorsal surface obliquely, a system of undulations is observed, which corresponds, though in a much more disguised form, with the condition in Scyllarus, the two extreme lateral pieces with the accom- panying spines being easily seen, and the small elevated centrum at the anterior part. In conclusion, an attempt has been made to show (1), That in some Macrurous Crustacea, and therefore probably in all, the telson is a true body segment with lateral appendages, which are modified by cohesion and adhesion. (2) That the body of the segment terminates posteriorly in front of the anus, and that the cohesion of the endopo- dites superiorly causes the anus to be ventral. ie i \Central Element { Endopodite Propod ite Telson, &e., of Scyllarus arctus. Page 356. Page 357. 96 NOTES ON AN OSTRICH LATELY LIVING 14, NOTES ON AN OSTRICH LATELY LIVING IN THE SOCIETY’S COLLECTION. (Written in con- junction with FRANK Darwin, B.A.)* A mate Ostrich (Struthio camelus) has been in the Society’s Gar- dens since April, 1869, and was quite healthy until last October, when its appetite began to fail, and it did not take kindly to its food from that time until its death on the 6th ult. In September last the keeper noticed on several occasions that after running about as it was accustomed to do in play, it turned giddy and apparently tripped, but never quite fell. For the last four months it had lost flesh gradually. Whenever any fresh food was offered it, it would take a little and then refuse any more, and would do thus, however many new things were ~ presented to it. It had suffered from diarrhoea more or less ever since October, the excrement having a yellowish-green colour. Latterly it had been nearly continually in the sitting position, and would stand very unwillingly. It also frequently rubbed its head and eyes with its foot, as if something was irritating it there. In the post-mortem examination very little structural disease was found; and the cause of death is more probably connected with the contents of the stomach rather than with any other agency. There was more than half a gallon of stones in the stomach: most of them were about the size of cob-nuts or peas; and they fully dilated the organ and pulled it down abnormally. Mixed up with these stones were numerous copper coins and pieces of coins in a much worn state. There were two pennies and fifteen halfpence; and very few showed the least trace of the stamp they had previously borne, and those only by an oblique light, the difference in density of the metal, produced by the stamping, having caused them to wear unevenly. Most of them were slightly curved, being meniscoid in form. They were all highly polished, and not in the least corroded. Many were in pairs, with a layer of softish green matter, about ;'5 of an inch thick, interposed. The chips of coins were very numerous, and of all sizes below that of the coins themselves. No silver was found, and nothing else except a glove-button and a nut, the latter being at the bottom of the sso- phagus. * “ Proceedings of the Zoological Society,” 1872, pp. 356-63. Read, March 5, 1872. IN THE SOCIETY’S COLLECTION. 97 All the contents of the stomach were of a green colour; and two small boluses of hay which it contained were tinged deeply with ' Four more coins, deeply corroded and greenish-black, were found in one of the intestinal ceca, together with a few stones. There were also a few stones in the other cacum; and the mucous membrane of both ceca was congested and unhealthy in appearance, which was not the case in the stomach to any extent. There were no symptoms of jaundice, which frequently accompa- nies copper poisoning. The liver appeared healthy, except that scat- tered abont were a few dense white lumps about the size of peas, mostly near the surface: it weighed 3 1b. 9 0z. No gall-bladder was present. The spleen was very small, and altogether weighed just under 2 oz. There was very little healthy tissue preserved, it mostly con- sisting of spheroidal dense masses of matter which were about the size of chestnuts, and by protruding beyond the general surface produced an appearance of knobs. These masses, on cutting through the capsule, separated entirely, and were then seen to be rough and altogether very like urinary calculi; they were of a fawn-colour. The organ was situated nearly in the middle line, just above the kidneys. The heart weighed 1 Ib. 7 oz., and gave origin to two carotid arteries, one from each main branch, which ran to the head, a dis- tance of about 3 feet 6 inches, side by side, in front of the cervical vertebre, in the groove formed by the anteriorly projecting processes of those bones; and they never showed any tendency to unite or Page 358. cross one another. They were thickly covered by the anterior cervical muscles, and sent off symmetrical branches.* Superficially on each side of the neck ran a vein with the pneumo- gastric nerve; but that on the left side was not bigger than a crow- quill, while that on the right had a diameter at the lower part of the neck of two-thirds of an inch. This condition is constant in many birds. This right (practically the only) jugular vein, after coursing abont half or a little more up the neck, sent two branches to the head, the second running in the middle line, just behind the trachea and in front of the cesophagus, the first being a direct continuation of - the main trunk. The intestinal canal was 34 feet long; and the two ceca, each * The presence of the two carotids in this bird, while there is only one in Rhea, would require that they should be far separated in Nitzsch’s classification of birds according to the number of these vessels—the Ostrich being in his first class, with a carotid from each main aortic branch, and the Rhea in the fourth class, with only the left developed. See Nitzsch’s “‘ Pterylography” (English edition), App. p. 171. H Page 359. 98 NOTES ON AN OSTRICH LATELY LIVING 2 feet long and arranged like a spirally twisted cone, were situated 11 feet from the pylorus, which is very different from their situation in most birds, as has been noticed by Owen. The diaphragm was well marked. It formed a partition which divided the thoracic cavity into two parts, one posterior and small containing the lungs, and the other anterior and large containing the heart and liver. It was a fibrous membrane, concave forwards, with a muscular attachment at either side to the ribs and intercostal tissues, which it joined in about the middle of their course. This muscular part was formed of transverse fibres in the middle and upper part of the chest, while the lower ones slanted slightly upwards as they coursed towards the median line. They were about 2 inches long, and formed a thin layer. The pleural cavity was closed above and below by the fibrous diaphragm becoming blended with the first and last ribs. The anterior thoracic cavity, which contained the pericardium- coated heart in its upper part, entirely independent of the pleural cavity, was divided into two by a dense fibrous membrane which sprang from two vertebral crura, much as the human diaphragm, and extended above the line to join the sternum, along the border which articulated with the ribs, leaving the heart entirely in front of it; its concavity was directed downwards and forwards; and: it was separated from the diaphragm proper by very large air-cells. The cesophagus also ran in the interval; but the aorta was included in the pleural cavity, being clearly seen through the membrane of the diaphragm, along the median line, before its removal. The liver was completely separated from the abdominal cavity by a fibrous membrane, so that when the included viscera had been re- moved it was not at all brought into view. The mesentery was very dense and strong, the vessels, especially the veins, being of large size. : Some further points in the anatomy of this bird are not without interest. There are three parietal abdominal muscles as usual, the muscular fibres of the external and internal being nearly parallel and transverse, while those of the intermediate one are longitudinal. They each send down a dense fascial attachment to the pubic bone; and a semilunar free margin between the ilium and the superior pubic crest appears closely allied to Poupart’s ligament, the anterior crural vessels and nerves going underneath it to enter the leg. It may be here men- tioned that the main vein of the thigh is the internal saphenous; but the main artery is the one that goes through the sciatic notch, there- fore the sciatic. These come into relation with one another in the loop for the biceps tendon at the knee, IN THE SOCIETY’S COLLECTION. 99 Exactly in the middle of the anterior border of the pubic portion of the innominate bone there is a small thin plate of osseous tissue which is connected with the pubis by strong fibrous bands, and which is continued anteriorly and superiorly by cartilage for some distance, when it becomes continuous with the tendons of the parietal abdo- minal muscles, being most connected with the external oblique. ~ Portions of the external surface of the left pubis and ischium of the Ostrich. The - small osseous plate (a) attached to the pubis is represented partly surrounded by cartilage. In dry skeletons a slight thickening of the anterior border of the pubic bone indicates the attachment of this ossification in most; but in one of the three skeletons in the British Museum this bone is an- chylosed on one side, and Mr. Gerrard has specimens in which both are still attached. A diagram of the Ostrich’s pelvis in Mr. Haughton’s paper also shows this bone anchylosed, though no mention is made of it in his paper. It would be extremely interesting to make out the homology of this small but perfectly independent ossification. Its relation to the muscles of the abdominal wall would favour the idea of its corre- Page 360. sponding to the marsupial bone of the Kangaroo and its allies; and if that is the case, the whole of the anterior prolongation of the Ostrich’s pubis would correspond to the small ridge of bone on either side of the superior margin of the symphysis pubis in the Mammalia. The obturator internus also arises from the superficial surface of this bone and its cartilage, as well as from the adjacent surface of the ischium and from the pubis, extending so far forward that the muscles of the opposite sides are only separated from one another by an inch or so at the symphysis pubis. H 2 Page 361, 100 NOTES ON AN OSTRICH LATELY LIVING Mr. Macalister, in his description of the myology of this bird, has omitted a few of the muscles, some of which from the head will be described, together with those of the leg, which Mr. Frank Darwin has allowed me to introduce in this communication, from his notes and the dissection of that limb in this individual specimen. Pterygoid.—F rom the inferior surface of the posterior part of the palate-bone, and from the process of bone which connects it with the main portion of it, this is fibrous—also from the whole of the inferior surface of the pterygoid bone, extending inwards almost to the basisphenoidal rostrum.. The fibres are all directed backwards, and are inserted in two ways :—the outer, and some of those from the palatine longitudinal process, into the anterior surface of the transverse ridge at the angle of the mandible, which posteriorly receives the insertion of the digastric muscle; the inner, and others from the palate-bone, into a fibrous band which runs from the side of the median Eustachian aperture and its cartilaginous continuation to the prominent ridge behind and internal to the dondyloid articular surface for the man- dible, thus forming an arch under which run the arteries and veins to the head. This second portion of the muscle acts partly as an opener of the Eustachian aperture, partly asa retractor of the slightly movable pterygoid and palatine bones. Quadrato-mandibular—From the whole of the longitudinal ridge which forms the superior internal portion of the quadrate bone, and from the surface of the bone external to it. The fibres are directed outwards and downwards to be inserted into the inner surface of the mandible, in front of the articulation, not extending to the inferior margin, nor forwards further than the optic foramen. Quadrato-cranial.—From the back of the orbit, below and behind the origin of the recti muscles and the exit of the nerve, from a surface bounded above by a semicircular line, and extending down in the space between the orbit and the quadrate bone. The fibres are directed outwards to the corresponding internal surface of the quad- rate bone, a slight ridge separating the superior ones from those of the quadrato-mandibular. Gastrocnemius consists of two enormous masses of muscle blending together at their origins round the proximal end of the tibia, and separating lower down into the gastrocnemius anticus, which laps round the anterior half, and the gastrocnemius posticus, which sur- rounds the back part of the tibial section of the limb. Gastrocnemius anticus arises partly from the tibia, partly by blending with gastroc- nemius posticus; the latter arises from the distal extremity of the femur, the tendon of the quadriceps extensor and patella, and from wee Ye ae | IN THE SOCIETY’S COLLECTION. 101 the tibia. At the tibio-tarsal joint the gastrocnemii form a sheath fitting into the trochlea of the tibia for the passage of the flexor ~ tendons of the toes; this is effected by the tendons becoming very | much thickened and semicartilaginous (especially gastrocnemius pos- ticus), and uniting with each other at their edges, the anterior element of the sheath being formed by gastrocnemius anticus, the posterior by gastrocnemius posticus. Just above the joint, gastrocnemius anticus sends off a slip which passes down in a special sheath along the outer surface of the contiguous heads of tibia and tarso-metatarsal bone, and is inserted into the tendon of flexor perforatus. Gastrocnemius anti- cus is inserted into the posterior surface of the tarso-metatarsal bone just below the tibio-tarsal joint. Gastrocnemius posticus is inserted into the external and internal lips of the posterior border of the tarso-metatarsal bone, forming a sheath for the passage of the flexor tendons; it subsequently forms, with a “sesamoid”’ cartilage presently to be described, a pulley for the same tendons at the tarso- _ phalangeal joint, and ends by blending with the fascia covering the sole of the foot. Mr. Macalister* describes the gastrocnemius as ending in one tendon only, which he says forms a sheath for the deeper tendons on the back of the metatarsus. The flexors of the toes are flexor magnus (perforatus), flexor per- forans, flexor externi digiti, flexor interosseus, flexor profundus. Flezor magnus arises (1) by a tendon from the upper part of the external surface of the outer condyle of the femur, the tendon winding over the knee, and then ending in the muscle; just before it does so, it receives the insertion of the rectus femoris (Cuvier and Meckel), (the pectineus of Owen); (2) from posterior surface of distal end of femur; (3) proximal end of tibia. The muscle ends in a broad tendon, which passes through the gastrocnemial sheath at the tibio- tarsal joint, and is here pierced by the tendon of flexor externi digiti. It passes down the tarso-metatarsal bone in the sheath formed by gastrocnemius posticus, receiving a tendinous slip, already described, from gastrocnemius anticus. At the tarso-phalangeal joint it passes through a sheath formed anteriorly by a “‘sesamoid” cartilage, posteriorly by the tendon of gastrocnemius posticus. This cartilage is ligamentously attached to the proximal end of the first phalanx of the internal digit, and to the synovial membrane of the tarso-pha- langeal joint ; it is deeply grooved posteriorly for the flexor tendons, and has two shallow grooves on its anterior surface, which fit on to the condyles of the tarso-metatarsal bone when the digits are ex- * Alexander Macalister, L.R.C.S.I., Demonstrator of Anatomy, Royal College of Surgeons, “On the Anatomy of the Ostrich (Struthio camelus),” “ Proceedings of the Royal Irish Academy,” IX. Part 1 (1865). Page 362. 102 NOTES ON AN OSTRICH LATELY LIVING tended; there is a smaller “‘sesamoid”’ cartilage for the external digit. A small muscle arises from the larger cartilage and by a few fibres from the smaller one, and is inserted into the anterior surface of the flexor profundus tendon. The flexor magnus is pierced, as it passes through the sesamoid sheath or pulley, by the tendons of flexor profundus and flexor perforans, and ends by dividing into two slips, which are inserted into the proximal end of the second phalanx of the internal digit. Mr. Macalister gives as the origin of the flexor magnus the deep pit above the condyles of the femur, the tendon of the rectus femoris, the external lateral ligaments, and the back of the fibula. In the specimen which I dissected, the tendon of the rectus femoris was much smaller than the tendinous head of the flexor profundus, and was in- serted into it, which is an arrangement differing very slightly from its usual insertion in birds, which is, I believe, into the fleshy part of flexor magnus (Owen, ‘‘ Anat. Vert.,”’ Vol. II, p. 107). The Rev. S. Haughton* describes the rectus as becoming provided with a second muscular belly (p. 53), which does not all describe its condition in my specimen. Heconsiders this “ digastric rectus femoris muscle” to be “‘the key to the explanation of the complicated museu- lar apparatus of the Ostrich’s leg” (p. 50). Speaking of it, he says, “it acts before the extensor muscles come into full play; it binds down the two patelle, braces up the heel-joint, and gives the signal for the m. gastrocnemio-soleus and other associated muscles to con- tract, and thus produces what may be regarded as one of the most striking phenomena in nature, viz., that the delicate bones and liga- ments of a bird’s leg, acted on by muscles equal to those influencing a horse’s hind leg, shall remain uninjured under the sudden action of forces the slightest error in the application of which would break to pieces the machinery on which they act.” This arrangement of the rectus, which Mr. Haughton considers so important, is only a well- developed form of what is found in most birds, and which Professor Owen says is used in perching, by flexing the toes when the knee is bent (loc. cit.) Mr. Macalister does not mention the muscle from the flexor profun- dus tendon to the sesamoid cartilage; but says that the flexor magnus sends a slip to it. The function of this muscle must be to keep the pulley-like sesamoid cartilage firmly in its place when the toes are extended preparatory to their flexion in the spring of the bird. * The Rey. S. Haughton, M.D., “‘ Notes on Animal Mechanics.—No. 3. On the Muscular Mechanism of the Leg of the Ostrich,” “‘ Proceedings of the Royal Academy of Dublin,” IX. Part 1 (1865). IN THE SOCIETY’S COLLECTION. 103 Flezor digiti interni arises from the outside of the tubercle of the tibia, and from the tendon of the quadriceps extensor; the tendon passes through the gastrocnemial sheath and the sesamoid pulley. It pierces the flexor magnus, receiving a tendinous slip from it, and then spreads ont and binds the tendon of the flexor profundus to the grooved under surface of the second phalanx of the internal digit, and is in- serted by two slips into the proximal end of the third phalanx. Flexor digiti externi arises from the posterior surface of the distal end of the femur, and from the tendinous head of flexor magnus. It passes through the gastrocnemial sheath, pierces the tendon of flexor Page 363. magnus, passes through the lesser sesamoid sheaths, and is inserted into each of the three proximal phalanges of the external toe. Flezor profundus arises by two heads—one from the posterior sur- face of distal end of femur, the other from the posterior surface of the upper half of the tibia and part of the fibula. The tendon of the external muscle passes at the tibio-tarsal joint through a canal in the tendon of gastrocnemius anticus. The inner tendon does not enter the gastrocnemial sheath till below the joint; it passes down the inner surface of the contiguous ends of the tibia and the tarso-metatarsal bone, bound down to them by a special aponeurotic sheath, and joins the outer tendon near the tarso-phalangeal joint. The common tendon - passes over the sesamoid pulley, piercing the flexor magnus tendon ; it is here much thickened and hardened, and fits into the grooved and thickened tendon of flexor perforans. It is inserted into the fourth phalanx of the internal toe, sending off a short strong slip to the third phalanx, and an elastic slip to the second phalanx, as well as a small but long slip to flex the fifth phalanx of the external digit. Mr. Macalister describes the tendon of the flexor profundus as being inserted only into the last phalanges of both digits; the insertion as it was in my specimen accords, I believe, with the usual condition of this tendon in birds. The flexor interosseus is a delicate and weak muscle, which con- sists of numerous very short oblique fibres arising from the posterior surface of the tarso-metatarsal bone, and inserted into an aponeurosis stretching the whole length of the muscle; this aponeurosis ends in a tendon which is inserted into the outer surface of the first phalanx of the external digit. Its action appears to be to abduct and flex the ‘ toe. | A small muscle which Mr. Macalister does not describe, but merely : mentions as probably representing the dorsal interosseus, arises from a small triangular space on the anterior surface of the distal end of the tarso-metatarsal bone, and is inserted into the capsular ligament of the tarso-phalangeal joint. There were some differences between the a ee —_— 104 NOTES ON AN OSTRICH, ETC. specimen which I dissected and that described by Mr. Macalister, in the precise origin of some of the muscles, which I have not thought to be worthy of note. The extensor communis digitorum presents no peculiarities ; the very small eatensor unguis mentioned by Mr. Macalister was present. THE MECHANISM OF THE GIZZARD IN BIRDS. 105 15. ON THE MECHANISM OF THE GIZZARD IN BIRDS.* Notwitustaxpine the fact that the external form and general Page 525. structure of the gizzard is known to almost every one, very little seems to have been. made out as to the means by which this organ is enabled to crush and render available for nutrition the hard grains taken as food. By most writers the gizzard is supposed to act as a grinding-mill, moving from side to side, assisted in its work by sharp-pointed stones which its owner swallows for the purpose. This was evidently the opinion of Hunter, though he seemed scarcely satisfied on the point when he found that there was no perceptible lateral movement of the muscular masses during digestion. Harvey gave a very good description of the action of the gizzard, as far as he knew it, in his description of the abdominal viscera of the common fowl (“On Generation,” Exercise vii); and Hunter is the only physiologist who seems to have worked at the subject since that time. The structure of the gizzard as a specialised organ is best seen in the Anserine birds; and that of the Goose will be now described. Externally it is circular when looked at from in front, oval from the side, and fusiform from above or below. The cesophagus enters it as a large infundibuliform tube, with the broader end downwards at its highest point; and the duodenum is continued ont of it behind Page 526. and above. The organ may be shown to consist of two lateral masses of muscle, with an oblong cavity between them, which opens above and below into two sacs, with muscular walls of nearly uniform thickness. The anterior superficial circular view presents the appearance of a central tendinous area, from which four lines radiate, nearly at right angles to one another, in an X-like manner. The upper and lower median areas between the corresponding limbs of the X are muscular and rounded at the margin, with the fibres directed to the central tendon. The lateral spaces are covered with glistening tendon, which at the edges shades into muscular fibres, not in this case curved, but straight and consequently squared off. The superior and inferior median portions are parts of the walls of * “Proceedings of the Zoological Society,” 1872, pp. 525-9. Read, April 16, 1872. 106 THE MECHANISM OF THE GIZZARD IN BIRDS. the corresponding cavities already mentioned; and the cesophagus enters the former at its inner angle, close to its junction with the right lateral mass ; the duodenum being behind. The lateral masses, with their tendinous coverings, are the mus- cular portions; and the cavity between them is just behind the central tendon. The epithelial lining of the whole organ is very dense, and is con- tinuous through the different cavities, terminating abruptly at the entrances to the cesophagus and the duodenum. Very shortly after the death of the bird it can be stripped off entire.* It is particularly dense where it covers the two lateral muscles, and generally forms a callous oval pad over each, which has to receive most of the force of the muscular walls as they act on the stones and food. The central tendons, one in front and the other behind, are very strong; and so are the fibres which radiate outwards from them; they are almost entirely connected with the lateral muscles. The lateral muscular masses have their fibres all tending forwards _ and backwards, each being inserted into both the front and back Page 527. tendinous expansion, the central being nearly straight and the lateral ones being curved slightly outwards in the middle of their course. The superior and inferior sacs are surrounded by muscular bands which bow over from front to back, being inserted into those parts of the margin of the central tendon to which they are opposite. By their contraction they reduce the size of the sacs and force any thing they contain between the lateral muscles, a considerable fold of the gizzard-lining, which acts as a kind of valve, preventing any stones entering the duodenum. The action of the lateral muscles can be best understood by observ- ing a horizontal section made through the middle of the gizzard. The section is fusiform and exhibits a central oblong cavity, short from side to side, bounded before and behind by the central tendons, and laterally by the triangular muscular masses. The accompanying figure and the above description show that in the gizzard there is no mechanism which could in any way produce any lateral movement of the one mass of muscles on the other; and it is difficult to conceive any epithelium, however horny and dense, that could resist the tearing-strain which would necessarily be associated with such movement, in addition to which several gizzards that have passed through my hands have been so loaded with fat or adherent to the abdominal walls, that any lateral movement must have been impossible in them. * This coat is considered by recent German authorities to be a secretion from the deep glands, not an epithelium. 107 Fig. 1. Horizontal section of the gizzard in a state of relaxation. Fig. 2. The same in a fully contracted state. The following explanation of the action of the gizzard as a simple crushing-organ seems to me much more in accordance with the known principles of animal mechanics. As is well known, muscular tissue, when it contracts, does not alter in volume, but gains in breadth what it loses in length during its action. Consequently when a large mass of short muscular fibres contracts it must alter its shape considerably, increasing greatly in breadth. This fact being borne in mind, the action of the gizzard is easily explained. The two enormous lateral muscles, with their fibres tending for- wards and backwards, when relaxed, have a large cavity between them, into which the seeds and stones are thrust by the simultaneous pyz¢ 52s. contraction of the superior and inferior muscular bags. Directly these have become fully contracted, the lateral muscles act; and by approximating the anterior and posterior tendons they become greatly expanded laterally. But this expansion can take place in one direction only—namely, towards the gizzard-cavity; for the anterior and posterior tendons being situated obliquely with regard to one another, and the contraction taking place through the whole mass, Page 529. 108 THE MECHANISM OF THE GIZZARD IN BIRDS. expansion can only occur towards the base of the triangle. The motion in this direction is furthered by the arrangement of some of the muscular fibres, as can be seen on close inspection of the section of the relaxed gizzard; for the dense horny pads above referred to are cupped on their attached surfaces, and the fibres run from one margin of this cup to the other, in an arched manner, as seen in the section. Those fibres just above the cup are arched also in the same way, and the epithelial margins of the cup are more yielding than elsewhere. Consequently, when the contraction occurs, the fibres straightening reduce the antero-posterior diameter of the cup and make the pad more convex towards the intermediate cavity, and push each towards its fellow, this action, combined with that of the other more marginal fibres, producing a most powerful compression of the contents. The great force exercised laterally by the contraction of a muscle can be well shown by tying a piece of tape round the middle of the arm proper, and then causing the biceps to contract forcibly, whereby the tape is broken. As remarked by most writers on the subject, every intermediate condition of muscularity of stomach may be found in birds, from the simple non-tendinous one of the Raptores and others to the most muscular of the Anserine birds. The degree of muscularity depends on the nature of the food which the bird obtains, as shown by Hunter’s experiment, in which he, by giving animal food to a duck (I believe),* caused a great diminution in the muscularity of its gizzard. The state of the bird as to health also influences the development of the muscular fibres, the heart and gizzard being very similarly affected by impaired nutrition. In the Gallinaceous and Passerine birds there is seldom a callous pad formed over the lateral muscles, the epithelium being generally plicated at right angles to the direction of the muscular fibres; and in them the organ seems to be a more simple squeezing-organ, though when rigor mortis occurs in a contracted gizzard it is seen that the muscular masses are convex on their opposed faces. From these remarks and what has been previously observed on the subject, the following summary statement may be made :— The gizzard is an organ which crushes, and so renders assimilable the harder portions of the food of birds. This food, having been previously macerated in the proventriculus or crop, is thrust between the lateral muscles (where it gets mixed with the small sharp stones it meets there) by the contraction of the superior and inferior gizzard- sacs—upon which these lateral muscles contract simultaneously ; and * [The bird experimented on was, in reality, a sea-gull. Cf. Everard Home, “ Lectures on Comparative Anatomy.” I. p. 271. 1814, Ep.] THE MECHANISM OF THE GIZZARD IN BIRDS. *109 their arrangement is such that all the force of their contraction is con- verted into a compressing force at right angles to their direction. This force, by tending forcibly to obliterate their included cavity, comminutes the more yielding of their contents and squeezes from between them the resulting chyme, which finds no difficulty in enter- ing the small orifice to the duodenum.* * Since writing the above, I find there is a peculiarity in the gizzard-pads in the swan and goose, which causes a slight up and down movement of the lateral mus- cular masses when in action. The lower end of one pad and the upper end of the other are much more strongly developed and are thicker than the rest ; this causes them to present a surface of contact one with the other, which is somewhat oblique with regard to the axis of the lateral muscles. Consequently, when these muscles come into play, the oblique surfaces tending to come into contact, the material to be crushed intervening, they, being opposed inclined planes, slide slightly on one another, the one mass rising while the other descends. During the diastole of the gizzard it resumes its former relations, and a reverse sliding occurs. Page 643. Page 644. 110 THE ANATOMY OF THE HUIA BIRD.. : 16. NOTES ON THE ANATOMY OF THE HUIA BIRD | (HETERALOCHA GOULD1).* A LivinG specimen of Heteralocha gouldi (Neomorpha gouldi, Gould, B. of Australia, IV. Pl. xix) was obtained by the Society on May 11th, 1870, as was announced by Mr. Sclater in the “ Proceedings,” 1870, p. 388. It died on the 28th of February, 1872, in a much emaciated condition, but without organic disease. The following notes relate to its anatomy, and may, I trust, assist in enabling its affinities to be more easily determined. Pterylosis—The arrangement of the feathers is completely passerine. The rhombic saddle of the spinal tract does not enclose any ephippial space, therein differing from the Crow’s and resembling the typical Starling’s. There are nineteen remiges, of which ten are on the hand ; they increase in size up to the fifth. The rectrices are twelve in number. The oil-gland is nude. Tongue.—Simple, horny, one-third the length of the beak. It forms a flat elongated triangle, slightly bifid at its apex, and a little prolonged backwards at its lateral borders, enclosing a curved line for the base, the concavity being backwards and carrying retroverted papillee. The mucous membrane of the palate extends forward as far as the middle of the tongue; that of the mandible goes a little further. At the angles of the mouth, just below the eyes, are two yellow oval cutaneous expansions, fixed in front and free at their borders else- where; they appear as if they were prolongations outwards of the mucous membrane of the angles of the mouth, which had been reflected backwards—they being continuous in front, round the margins, with the mucous membrane. Syring.—As in Corvus and most of the Old-World Passerines. Intestines —The gizzard is well developed. The intestines are 16 inches long, with the bile-ducts 24 inches from the gizzard. The ceca are one inch from the cloaca and + inch long, being cylindrical. Arterial System.—There is one carotid artery, the left. Foot.—The hind toe is slightly longer than the middle anterior toe. In arrangement the tarsal scutes are similar to those of Corvus and most Passerines. Their colour is blue-black. Skull.—The palate (Fig. 2) is strictly sgithognathous; that is, * “Proceedings of the Zoological Society,” 1872, pp. 643-7. Read, May 21, 1872. THE ANATOMY OF THE HUIA BIRD. 111 the vomer is truncate in front abruptly, and cleft behind; the postero- external angles of the palatines are produced; the maxillo-palatines are slender, and approach towards but do not unite with one another, Fig. 1, Skull of Heteralocha gouldi, lateral view. Fig. 2. Skull of Heteralocha gouldi, inferior view. Fig. 3. Skull of Heteralocha gouldi, posterior view: d_f, digastric fossa. Fig. 4. Skull of Corvus frugilegus, posterior view: d.f, digastric fossa. Page 645. Page 646. 112 THE ANATOMY OF THE HUIA BIRD. nor with the vomer, which they partly embrace. There is no ossifica- tion in the nasal septum anterior to the vomer. The whole cranial configuration (Fig. 1) closely resembles that of Sturnus ; but the mandible, instead of being bent upwards, is straight. Like it, the palatines are narrow and approximate; the antero-internal angles of the posterior portions of those bones are reduced and rounded off, as is sometimes the case with Stwrnus (Mus. Roy. Col. Surg. No. 1539, Ost. Coll.). The vomer is completely truncated in front, and is not prolonged forwards at its external angles, as in Corvus and its allies. - The zygoma is not so slender as in Sturnus; but the curves are similar. The articular surfaces on the quadrate bone for the mandible are proportionally very large. The anterior extremities of the pterygoid bones articulate with the sphenoidal rostrum much as in Corvus, meeting in the middle line behind the posterior extremities of the palatines for a short distance. The maxillo-palatines in their approximate portions are shorter from before backwards than in Stwrnus, and much resemble those of Corvus. The antero-inferior processes of the orbit are large and spongy ; they almost touch the zygoma. But the most characteristic portion of the skull of Heteralocha is the occipital region; and in this it presents a great exaggeration of the peculiarities of Stwrnus and its allies. In Corvus (Fig. 4) and most Passerines the digastric muscles occupy a narrow space intervening between the auditory meatus and the mass of occipital muscles, not extending so high up the skull as the latter. The occipital ridge encloses a space elongated from side to side and of but little depth. In Sturnus the digastrics are much broader, and they narrow the occipital space; they also extend up the skull to so great an extent that they nearly meet in the middle line above the origin of the biventres cervicis muscles; but in Heteralocha (Fig. 3) they are of still greater size, and meeting above the middle line they form a strong ridge, which extends for some distance into the parietal region verti- cally. This peculiar development of these muscles produces a corre- sponding change in the shape of the space enclosed by the occipital ridge. In Heteralocha it is almost circular, and it extends some way above the foramen magnum. In Stwrnus there is an approximation to this condition. A vertical parieto-occipital ridge in many other birds closely re- sembles that of Heteralocha; but it is the median limit of the temporal fossa in most. Correlated with this extensive digastric origin is a large surface THE ANATOMY OF THE HUIA BIRD. 113 for its insertion. The angle of the mandible (see Fig. 1, p. 647) is prolonged directly backwards for this purpose, in a manner unique among Passerine birds, but well seen in the Anatide: In Sturnus the angle of the mandible is slightly prolonged backwards for a similar per pose. 4 In comparing the skulls of others of the Sturnide the following is a graduated series, based on the development of the digastric fossz in those birds I have had the opportunity of examining, commencing with Heteralocha and ending with Corvus. Heteralocha. Quiscalus. Sturnella. Cassicus. Sturnus. Acridotheres. Icterus. Pastor griseus. Pastor jalla. Molothrus. Gracupica. Corvus. The palates in most of these birds were not in a fit condition for study ; and, as will be clearly seen, geographical range has not been attended to. In the sternum, Heteralocha differs in no important point from _ Sturnus, except that the posterior notches tend to be converted into foramina, as observed by Mr. Eyton in his ‘‘ Osteologia Avium.” The following muscles were dissected, and found to agree precisely Page 647. with the corresponding ones in a Rook. Pectoralis major. — Sartorius. Pectoralis minor. Semimembranosus. Coraco-brachialis longus. Semitendinosus. Coraco-brachialis brevis. Adductor magnus. Tensor patagii longus. Biceps. Tensor patagii brevis. Femoro-caudal. Tn conclusion, it may be stated that the anatomy of Heteralocha shows clearly that it is truly Passerine, and not related to Upupa, as was previously supposed by most authors.* When examined more in detail, its relation to the Stwurnide is found to be very intimate, and its structure is clearly not closely allied to that of the Corvide. In its relation to Sturnus it seems to present an exaggeration of the peculiarities of that bird, which would place it at the head of the family. * Mr. G. R. Gray has placed Heteralocha in the Sturnide is his “ Hand-list of Birds.” Page 787. 114 THE TONGUE OF THE PSITTACINE GENUS NESTOR. 17. NOTE ON THE TONGUE OF THE PSITTACINE GENUS NESTOR* On the death of a specimen of Nestor hypopolius in the Society’s Gardens, a short time ago, Mr. Sclater kindly directed my attention to the peculiarity of its tongue, and referred me to Dr. Finsch’s work on the Parrots, where Nestor is placed among the Trichoglossinw, though the author states that he is not possessed of any very precise informa- tion on the subject. Mr. Gould, in his “ Birds of Australia ” (vol. 5, plate VI), partly describes the tongue of this bird, and shows that it is not that of a Lory; but he has omitted to note its chief peculiarity. Dr. Buller, in the recently published first part of his “ Birds of New Zealand,” has also described the tongue quite correctly, though not much in detail—but nevertheless places Nestor close to the Lories, mentioning that this affinity was first shown gf MM. Blanchard and Pelzeln. As, however, the tongue of Nestor does not in reality resemble that of the Trichoglossi at all, it may be of interest to describe it more fully. As far as I have had opportunity of observing, in all Parrots the fleshy tongue ends anteriorly by a dilated portion, supported on a narrower neck. This tip is much like the end of a human finger, as mentioned by most observers: and its function is similar also; for it is employed by the bird as a third prehensile organ in connection with the upper and lower beak, any solid substance being held by the tongue and upper beak, while the mandible is freed to give another hite. Continuing the simile of the finger, the tip is directed forwards with the nail-like portion downwards, the part corresponding to the free edge of the nail ene along the lower margin of the anterior rounded surface. This unguis, or nail-like portion, appears to me further to re- semble a nail in that its anterior edge is not quite regular and is free, while the posterior margin is continuous with the neighbouring epithelium, which is almost enough to show that it grows forwards, and is worn down, as is a nail, by constant contact with foreign sub- stances. * “Proceedings of the Zoological Society,’ 1872, pp. 787-9. Read, June 18, 1872. THE TONGUE OF THE PSITTACINE GENUS NESTOR. 115 In the typical Parrots this unguis of the tongue is broader than long, horny in texture, semicylindroid, with its lateral margins extend- ing up the sides of the organ and encroaching on the borders of the superior surface for a short distance; not imbedded at the sides as is anail. Its anterior border is nearly straight. In the Trichoglossi this horny plate is also present, and is similarly Page 788. constructed; but on the superior surface of the tongue, between the lateral edges of the unguis, in the part which in others is covered by a smooth longitudinally plicated epithelium, there is an arrangement of retroverted papille forming a spinous covering; and their mechanism is such that when the tongue is protruded beyond the mouth to grasp Fig. 1. Head of Lorius tibialis, showing the bird stretching out its bill for food, in which case, the tongue being protruded, the spines covering the superior surface of its apex are directed forwards instead of being recurved and inconspicuous, which is the case when it is at rest. Figs. 2 and 3. Inferior and superior views of the tongue of Nestor hypopolius, 4 showing the fringe of hairs which springs from the anterior border of the margins, , and which extends forwards beyond the tip. _ Figs. 4 and 5. Inferior and superior views of the tongue of Stringops habro- By ptilus, which is like that of the typical Parrots. 12 Page 789. 116 THE TONGUE OF THE PSITTACINE GENUS NESTOR. any object, the papille stand upright or are even directed somewhat forward. : In Nestor there are no papille of this description, but the tongue is here, as Dr. Buller says, “ soft, rounded on the edges, with a broad central groove,” and it is as smooth as in other Parrots. Therefore the Ka-Ka Parrot cannot in this point be said to approach the Tri- choglossi (badly so called). The peculiarity of the tongue of Nestor consists in the fact that the anterior edge of the unguis, always free (though for a very short dis- tance) and jagged, as mentioned above, in the other birds of the class, is here prolonged forwards, beyond the tip of the tongue, for about 15 inch as a delicate fringe of hairs, with a crescentic contour. This fringe seems to result from the breaking up into fibres of the forward- growing plate, which is always marked by longitudinal striations, clearest anteriorly, the result of unequal density and translucency of the tissue composing it, though on making a cross section I was not able to find any of the longitudinal papillary ridges which are present - in the human nail and which the striation led me to expect. The unguis is also longer than broad, and very narrow considering the size of the bird, as is also the whole tongue, though the length is greater than in others of the class. In the living bird the mouth is moist, as in the Lories, and not, as in the Cockatoos and others, dry and scaly. From these considerations, and a comparison of the accompanying drawings of the tongues of Stringops, Nestor, and Trichoglossus, it is evident that the structure of this organ would lead to the placing of Nestor among the typical Parrots, though an aberrant one, and not with the Trichoglossine ; and other points in its anatomy favour this conclusion. THE CRANIAL PECULIARITIES OF THE WOODPECKERS. 117 18. NOTE ON SOME OF THE CRANIAL PECU- LIARITIES OF THE WOODPECKERS.* Constperinc the method adopted by the Woodpeckers for obtaining page 357. their food, it is hardly surprising that they possess cranial features - peculiar to themselves; for it is scarcely conceivable that the head, the most delicately constructed portion of the body, should be em- ployed as a powerful hammer or axe whose stroke can be heard at a considerable distance, without some modifications in structure which would assist in increasing its efficiency for the purpose. Accordingly, we find that the bones are thicker and stronger than Page 358. in most birds, and there is only a slight movement possible of the upper jaw on the head proper. The interorbital septum is thick and nearly complete, supporting a median protrusion on the front of the skull, which is so considerable as to throw the free extremities of the hyoid bones one side or the other, thus causing the skull to be slightly unsymmetrical. Further, the axis of the upper beak is peculiarly low, being continuous with that of the basicranium ; and this results from the lowness in position of the points of junction of the superior processes of the premaxille with the frontal region of the skull, which renders the angle between the beak and skull less obtuse than is generally the case. In those birds in which there is considerable hinge-motion of the upper beak on the head, as in the Parrots, the basisphenoid rostrum is generally long and of uniform thickness for some distance, and the conjoined palatine bones, with the vomer between them and the pterygoids articulated behind, form a longitudinal flange along the upper service of the median junction, which runs backwards and forwards on the rail formed by the basisphenoid rostrum during the movements of the beak. In the Woodpeckers any considerable articulation of this kind would reduce the value of the head as an axe; consequently the posterior ends of the palatine bones are not well developed, and they scarcely ‘unite in the middle line, while further forwards the vomer is not seen in the maxillo-palatine region, and these latter bones also are only slightly developed. A similar tendency among Passeriniform birds to the reduction of the vomer in * “Ibis,” 1872, pp. 357-60. October 1. 118 THE CRANIAL PECULIARITIES OF THE WOODPECKERS. front is found, combined with a complete absence of the maxillo- palatines, in Menwra.* Professor Huxley in his paper “On the Classification of Birds,’ has entered into considerable detail respecting the Woodpecker’s palate, and from not finding a vomer present, and observing the peculiar longitudinal bony spicula connected with the inner edges of the palatine bones, opposite to and behind the fenestree they assist to Page 359. enclose, is led to think that these spicule are the rudiments of the vomer, which has not ossified across the middle line. But in carefully prepared skulls they look much more like the inner edges of the imperfectly ossified palatines, as they are connected completely with them at both ends. Further, in most of the specimens of Gecinus viridis and its allies that I have had the opportunity of examining, I have found a median bone, situated between the palatines, and sup- ported like a vomer on the basisphenoid rostrum, at the anterior ends of its broader portion. This bone is small, and shaped very much like a spear-head with the tip directed forwards, whilst posteriorly it gradually becomes fibrous and tends to bifurcate, but not in the ossified part. It does not extend backwards quite so far as the pterygo-palatine articulation. The accompanying sketch will enable its shape and position to be more clearly perceived. Palate of Gecinus viridis, showing the vomer 2, between the palatines Pl. The pterygoids are marked Pt, and the spine of the basicranium a. * Since the above has been in print, I find that the maxillo-palatines are not absent in Menwra, but are long and slender, differing somewhat from the ordinary Passerine type, but separate from one another and from the vomer. + “ Proceedings of the Zoological Society,” 1867, pp. 415-472. THE CRANIAL PECULIARITIES OF THE WOODPECKERS. 119 Though this bone is situated rather further back than the vomer in most birds, yet it is found similarly placed in some, as in Megalema, which by the way has the anterior termination of its vomer truncated in front, and produced forwards at the corners, as in the Crow, though in the former bird these processes articulate, and do not anchylose with the posterior ends of the palatine plates of the maxillo- palatines. On cutting the palatine bones of Gecinus from the anterior part of Page 360. the skull, and disarticulating them from the pterygoids, the bone which I suppose to be the vomer comes away with the palatines, as would be expected were such the case. The absence of truncation in the vomer of the Woodpeckers tends by itself to remove them from a close relationship with the Passerine birds; but, as I before remarked, this peculiarity may depend on their special habits. There is, however, in the shape of the pterygoid bones a character which tends to bring them together again. In Passerine birds the pterygoids extend forwards for a considerable distance in front of the point of contact or articulation with the pala- tines. These anterior processes are vertically expanded and in contact with the rostrum, and probably sometimes with the crura of the vomer; they are situated above (that is, deeper than) the posterior _ internal angles of the palatines, and therefore are not seen while ~ looking at the surface of the palate, but only on aside view. In the Woodpeckers and other birds related to them these processes are also present, but they are absent in most others, though the Anserine birds possess them. In the Woodpeckers also there is a very peculiar anteriorly directed process arising from the upper part of the middle of the body of the pterygoid bone, which is quite independent of the one above described. Page 821. Page 822. 120 THE PLACENTA OF THE HIPPOPOTAMUS. 19. NOTE ON THE PLACENTA OF THE HIPPO- POTAMUS.* Nor knowing of any description of the placenta of Hippopotamus amphibius, I think it desirable to record the condition of that obtained after the birth of the calf, which occurred on the 5th of this month. The placenta is a long cylindrical bag, 33 feet from end to end and 14 feet across. There is only one aperture; and that is not more than a foot long, and is situated at one of the ends. The other end is rounded, and quite complete. It is evident that the whole viscus is much the shape of the enclosed foetus, and must have closely covered it. The end at which the rupture had occurred, that is the one situated at the os uteri, is a little constricted, as may be inferred from the above statement of its diameter. The umbilical cord is attached to the placenta at one of the sides, about halfway between the two ends. It is 14 feet long, and ragged at its free extremity. It is 14 inches in diameter in the middle, and gets larger as it approaches its attachment, near which there are many spherical bodies, as big as peas and yellow in colour, supported on short amniotic pedicles. The outer surface of the whole viscus is covered uniformly with villi of a bright red colour; and there is no reduction of their number, nor in their size, at the cecal end at all. At the lacerated extremity, close to the rupture, they are paler and more scattered. The walls of the viscus are of uniform thickness, except for a few inches round the ~ point of attachment of the cord, where the vessels commence to diffuse themselves. When received, the whole sac was turned inside out; and this was probably the result of the gradual contraction of the uterus from fundus to orifice. It may be remarked that for a few days after the birth of the calf, the mother had a considerable prolapse of the vagina, which gradually diminished, and is now very slight. * “Proceedings of the Zoological Society,” 1872, pp. 821-2. Read, Nov. 19, 1872. THE ORDER DINOCERATA. ; 121 20. ON THE ORDER DINOCERATA (Marsx).* Ir is very seldom that specimens are now obtained, either fossil or Page 267. recent, of mammalian forms that have, provisionally at least, to be placed in a new order by themselves, on account of the presence of peculiarities hitherto quite unknown and unexpected. Such, how- ever, is the case with a large series of fossils which have been, within the last two years, obtained from the Eocene deposits of Wyoming in North America. Our chief source of information respecting these remains consists of a series of papers by Professor O. C. Marsh, of Yale College, abstracts of which have recently appeared in “The American Journal of Science,’ and in “The American Naturalist.” Professor E. D. Cope has also published several papers on the subject, and some excellent photographs of one of the most important forms sent by him to this country it has been my good fortune to see. Professor Leidy has also named one of the genera. Most of my information is obtained from the papers by Professor Marsh on the Order Dinocerata, together with the critical - remarks made by him on Professor Cope’s descriptions, which I have also by me. Dinoceras mirabilis is the name given by Marsh to a huge ungulate animal, of which there is an almost complete skeleton in the museum of Yale College. Its size must have been very nearly that of a full- grown elephant, as the length of the skull of a closely allied genus was a little more than a yard. The skull was peculiarly long and narrow, and supported three pairs of horn cores in rows, on its superior surface. The anterior pair were situated on the anterior ends of the nasal bones; they were short, conical, and directed nearly straight upwards. The median pair were conical prolongations upwards from the maxillaries; they were longer and more cylindrical than those on the nasals, and the fangs of the huge canine teeth entered their bases. The posterior pair, the largest, were very pecu- liar, being extensions upwards from near the middle of large lateral longitudinal crests, which were formed by the occipital, parietals, and Page 268. frontal bones. The dentition was peculiar; the upper incisors were deficient, and the premaxille consequently small. The upper canines formed huge downwardly directed tusks, nearly straight, but directed somewhat backwards. These, after a gap equal to their * “Journal of Anatomy and Physiology,” VII. pp. 267-270. June, 1873. oS a 122 THE ORDER DINOCERATA. breadth, were followed by six small molar teeth. Each of these were composed of two well developed, nearly straight, transverse ridges, which joined at the inner border of the tooth, and diverged as they Skull of Dinoceras mirabilis, copied from Professor Marsh’s figure, and kindly lent by the Proprietors of “ Nature.” ran outwards, so forming a simple >-shaped pattern, that continued well marked till the tooth was much worn. The posterior of the two ridges was quite transverse, and the anterior was slightly concave backwards, and ran obliquely forwards and outwards. These teeth Page 269. diminished in size from before backwards gradually, and there were no intervals between them. All that I can find about the lower jaw is that it was slender, and its tusks small. The orbit was not separated from the temporal fossa, which latter was large, extending up the outer side of the lateral parieto-occipital crest. The malar completed the anterior portion of the zygomatic arch; the lachrymal was large, forming the anterior border of the orbit; a large oval foramen perforated its facial surface. The squa- mosal sent down a large post-glenoid process. The nasals were massive, and greatly prolonged forwards, at the tip carrying the anterior horn-cores; they also sent slight processes up the inner faces of the maxillary horn-cores. The premaxillaries were peculiar in that they almost enclosed the anterior nares; they united posteriorly with the maxillaries just in front of the canines, and then divided into two branches, one of which, the lower, corresponding to the premaxillary of the Rhinoceros, ran forwards free; the other, closely uniting with THE ORDER DINOCERATA. 123 the adjoining nasal, went upwards to strengthen the support of the horn-cores. The extremities were tetradactylate, and in the pes the astragalus articulated with the cuboid as well as with the scaphoid bone. The humerus was short and massive, with the great tuberosity not rising above the articular head, and the condylar ridge of the distal end not continued up the shaft. The radius, which was free, was not so oblique as in the Elephant. The femur had no pit on its head for the ligamentum teres, and there was not any third trochanter. The phalanges were short, as in the Elephant. The number of the vertebre are not noted by Marsh, so that I cannot say how many there were in the dorso-lumbar region, a point of great importance. There were four in the sacrum, the last being small and supporting a slender tail. The ribs had rudimentary uncinate processes. The name Tinoceras has been given by Marsh to a very closely allied species, which differs from Dinoceras in having the anterior or nasal horn-cores compressed on the top, larger, and projecting more forward; the maxillary horn-cores are also proportionately longer, more cylindrical, and directed slightly forward. The photographs of + _ this genus above referred to show very clearly that the palate was ~ completed opposite the posterior molars by the palatine bones. The differences between Tinoceras and Dinoceras seem to be scarcely generic. Another closely allied genus, of which I have seen no description, is the Uintatherium of Leidy. The name Hobasileus, introduced by Pro- Page 270. fessor Cope, is a synonym of Tinoceras, and being of later intro- duction, must be sunk. From the facts given above, Professor Marsh is led to placing these undoubtedly peculiar animalia in an order different from any yet established, intermediate between the Proboscidia and the Perisso- dactylata. To me it seems much more probable that they belong to the Ungulata proper, and that no separate order is necessary for their reception. In the characters I have given, there are none which show them to have any true Proboscidian affinities, whilst there are several which seem to indicate that they belong to a family of the Artiodacty- lata, and not to the Perissodactylata. Among the reasons in favour of Dinoceras and its allies being Artiodactylate are the following :— The astragalus has a well-developed cuboid facet. . The palate is complete between the posterior molars. . There is no third trochanter to the femur. The premaxille are edentulous. The anterior premolar is not developed. gr 9 PO pt Page 33. Page 36. 124 THE NASAL BONES OF BIRDS, 21. ON THE VALUE IN CLASSIFICATION OF A PECU- LIARITY IN THE ANTERIOR MARGIN OF THE NASAL BONES IN CERTAIN BIRDS.* Since commencing the study of the anatomy of birds, it has always appeared to me that two distinct types of nasal bones can be dis- tinguished among them without difficulty—and that if those which present the abnormal characters are considered separately, they pre- sent other features in common which justify their being placed in the same class, and their entire separation from those which present the less modified arrangement. In most birds the anterior margin of the nasal bone is concave, with the two cornua directed forwards—one along the outer edge of the nasal splint of the premaxilla, to form the inner margin of the osseous external nares, whilst the other, which is free, descends as part of the external boundary of the same aperture in connection with the ascending process of the maxilla, which it joins. These two processes become continuous behind with the body of the bone, and with one another, there being no interruption of any kind between them. Such a condition is found in its simplest form in Otis and the Galline proper; and birds possessing the bone so constructed may be termed holorhinal: in them a transverse straight line, drawn on the skull from the most backward point of the external narial aperture of one side to that of the other, always passes in front of the posterior terminations of the nasal processes of the preemaxille. But several birds present a very different condition. In Grus, for example, the posterior contour of the osseous external nares, instead of being rounded, as in holorhinal birds, is apparently formed by the divergence of two straight bars of bone, which enclose an angular ‘space between them. These two processes evidently correspond to the two anteriorly directed cornua of the holorhinal skull described above ; but they appear in many cases to be so different in density, and the outer one joins the body of the bone so abruptly, that it seems at first sight to be an independent ossification; however, I have no reason to believe that such is the case. As in holorhinal birds, so in those under consideration, which may be termed schizorhinal, the internal process of the nasal bone runs forwards along the outer border of the * “ Proceedings of the Zoological Society,” 1873, pp. 83-8. Read, Jan. 7, 1873. THE NASAL BONES OF BIRDS. 125 Skulls of Schizorhinal Birds. Page 34. 3 1. Alcea impennis. 5. Parra (Hydralector) cristata. A 2. Larus argentatus. 6. Arctica alle. 3. Numenius arquatus. 7. Pedionomus torquatus. 4. Columba livia. 126 THE NASAL BONES OF BIRDS. Page 85. Skulls of Holorhinal Birds. 4 Ww f # Sy ® vw! He ) 8. Otis tarda. 10. Daption capensis. 9. Gallus domesticus. 11. Coccothraustes vulgaris. THE NASAL BONES OF BIRDS. 127 nasal process of the premaxilla, and the outer descends free to join the maxilla. In these birds there is. considerable variation in the manner in which the almost detached outer of the two nasal processes joins the body of the bone. In Numenius, Hematopus, and many of the Limicolz they proceed directly upwards and expand, becoming slightly fanned out where they join the rest of the bone by a straight transverse line. In Ibis and Grus they are of uniform size from end to end, whilst in the Auks, and to a less degree in the Gulls, at its _ origin the process is slightly curved, being directed outwards for a short distance, and after that straight downwards and forwards. In most schizorhinal birds, a transverse line joming the extreme posterior point of one external nasal aperture to the similar one of the opposite side is situated behind the posterior ends of the nasal processes of the premaxilla; but in some of the short-beaked broad- mouthed species of the class it is situated in front of them. Such is the case in Pterocles and Syrrhaptes ; and this peculiarity renders it at first sight uncertain whether they are schizorhinal at all; but as every intermediate condition may be found between the strictly schizorhinal skull of the Columbide proper, and the very similar but less strongly marked skull of Pterdcles, there is no real reason to doubt that the modification only depends on the great breadth of beak in the latter - bird. The curious development of the superficial nasal turbinal bone of Pterocles is also a Columbine character, as is also the great length of the inner of the two nasal processes, which, in a manner quite un- like that of the Galline, extends on each side for a long way forwards under the premaxillary nasal splint. Subjoined is a list, alphabetically arranged, of the genera in which I have observed the schizorhinal arrangement :— ScHIZORHINAL Birps. Alea. Grus. Rhinochetus. Anous. Hematopus. Rhynchops. Anthropoides. Ibis. Rissa. Arctica. Larus. Sarciophorus. Cataractes. Lestris. Scolopaz. Charadrius. Limosa. Sterna. Chionis. Machetes. Totanus. Dromas. Numenius. Tringa. Eurypyg. Parra. Turniz. Fratercula. Platalea. Uria. Gallinago. Pierocles. Vamellus. Glareola. Recurvirostra. And all the Columbidz proper. Page 37. 128 THE NASAL BONES OF BIRDS. From the above list it is evident that nearly all the schizorhinal birds are included among Professor Huxley’s Schizognathe. Going farther into detail, they may be said to embrace all the Charadrio- morphs, with the exception of Hdicnemus (I have not seen Cursorius). Among the Geranomorphe, they comprise the Gruide, together with Rhinochetus and Ewurypyga, but not the Rallide (from which family Parra should be removed to the Charadriomorphe), nor Psophia, Otis, and Oariama. Among the Cecomorphe they include the Laride and Alcides, but not the Procellariide nor the Colymbide. Among the Speniscomorphe none are schizorhinal. Among the Alectoromorphex Turniz and the Pteroclide are so; and the Peristeromorphe are all schizorhinal. In the other main divisions, the Desmognathez and the Aigithognathe, only the Hemiglottides of Nitzsch, belonging to the Pelargomorphe, are schizorhinal. The linking together of the Plovers and the Gulls by any osteolo- gical feature has long been a desideratum, as Professors Newton and Huxley have remarked ;* and the facts brought forward by the latter have greatly assisted in this respect. But Professor Huxley’s classifi- cation does too much; it places the Petrels nearer to the Gulls than the latter to the Plovers, and it includes the Rails in the same cate- ~ gory as the Cranes, which is more than collateral evidence justifies. If the nasal bones have the significance in classification which I would put upon them, and their conformation be employed in dividing up the schizognathous birds (with which, notwithstanding their desmog- nathism, Platalea and Ibis must be placed), a result is arrived at which pterylosis and internal anatomy greatly tend to justify. The following table represents my idea of such an arrangement, though I do not wish to give my sanction to the naturalness of the non-schizorhinal schizognathous group, which I believe to be open to criticism :— Scuizoanarnous Brrps (Huxley) + Hemiglottides (Nitzsch). Schizorhinal Birds. . Columbe, Pteroclide, and Turnicide. . Limicole (excluding Gdicnemus, and including Parra). . Laride. Gruide. Eurypygide and Rhinochetide. . Hemiglottides. . Alcides. SIO OP De * “This,” 1868, pp. 92 and 360, THE NASAL BONES OF BIRDS. 129 iediphinal Birds: Page 38. 1. Impennes. 2. Procellariide. . Colymbide. Galline (excl. Pterocles and Turnizx). Rallide (excl. Parra). Otide (incl. Edicnemus). Cariamide. Psophiide. Opisthocomide. . Podicipide. In his paper “On the Osteology of the Kagu,” Mr. Parker, in speaking of the nasal bone, says, “‘this part of the face is thoroughly Gruine in both the Eurypyga and the Kagu; the long open nasal fossa, so sharp above at the bifurcation of the nasals, gives a character to the face common to large groups of Gralle and Palmipeds.” Other- wise he does not employ this character in classification, as is evident when it is seen that he places the Kagu close to Psophia and the Rails, which are holorhinal birds. It may be mentioned that the external nasal process of the nasal bone is weak or obsolete in the Struthious birds. SOON A op pay Page 92. 130 THE VISCERAL ANATOMY 22, ON THE VISCERAL ANATOMY OF THE SUMATRAN. RHINOCEROS (CERATORHINUS SUMATRENSIS).* TE death on September 21st, 1872, of the only English specimen of the Sumatran Rhinoceros has afforded me an opportunity of deter- mining many points in its anatomy previously unknown; and Pro- fessor Owen’s excellent memoir on Rhinoceros indicus, in the fourth volume of the Society’s “‘ Transactions,” has made it possible to com- pare the details of structure in the two species. The differences in the shape of the stomach, and the character of the mucous membrane of the small intestine, together with the pecu- liavities of the skin, including the presence of a second horn, the absence of a gland behind the foot, and the smallness of the folds, which cannot accurately be termed shields, appear to me quite to justify the separation, into a distinct genus, of the Sumatran Rhi- noceros from its Indian ally, as has been done by Dr. Gray from a study of its osteology only. The specimen upon which these observations are based is said to have been captured in Malacca: it is an aged female: its skin is of a dark slate-colour, and is covered thinly with black hairs, which are more than an inch long, situated mostly on the middle line of the back and on the outer sides of the limbs. Its length from the tip of the nose to the base of the tail is 964 inches. The tail is 22 inches long; from its base to the transverse shoulder-fold is 44 inches; and from the latter to the occipital crest is 22 inches. The ears are lined, and not fringed (as are those of the Ceratorhinus lasiotis) with black hairs. No traces could be found at the back of the feet of the glands described by Professor Owen in the Indian Rhinoceros. The skull, the only part of the skeleton which I have examined, is 21% inches from the tip of the nasal bones to the middle of the occipital crest, following its longitudinal direction. From one lachry- mal tubercle over the head to that of the opposite side is 8 inches. The conjoined nasal bones in their broadest part are 6% inches across from their lower margins over the insertion of the anterior horn. The lower incisors and the first premolars are lost; Professor * “Proceedings of the Zoological Society,” 1873, pp. 92-104. Read, Jan. 21, 1873. + See for an account of its history Mr. Sclater’s notes, “ Proceedings of the Zoological Society,” 1872, p. 494. OF THE SUMATRAN RHINOCEROS. 131 Flower informs me that a specimen in the Museum at Brussels has also lost its lower incisors. The premaxillary bones are anchylosed to the maxillaries, a condition I have not found in any other specimen, and which is probably dependent on the loss of the lower cutting-teeth. Including the present one, I have seen eight skulls of Asiatic two- horned Rhinoceroses (Ceratorhinus)—four in the College of Surgeons’ Museum, two in the British Museum, and one at the Museum at Cambridge. The present specimen agrees very closely with that at the last-named place, and with No. 1461 a, adult, from Pegu, in the British Museum. It being that of an aged individual, comparison with most of the others referred to is more difficult, as they are Page 93. nearly all immature. The skeleton mounted at the College of Surgeons, No. 2933, obtained by Sir S. Raffles from Sumatra, is aged also; but there are points in which it differs materially from the present specimen. It is of slighter build, and the nasal bones are narrower, A much larger skull, not quite adult, in the Museum of the Col- lege of Surgeons, No. 2935, stated to be that of a “male Sumatran Rhinoceros,” and presented by Sir S. Raffles, is evidently from a larger animal, and agrees also with Ceratorhinus lasiotis in being pro- _portionately broader in the parietal regions. In this skull also the " posterior of the submental foramina is situated in front of the second premolar, while in all the others (except No. 2936, R.C.S., which is young, but peculiarly massive) it is situated, when present, behind that tooth. Subjoined is a table giving a few of the measurements in the skulls above referred to :— a Zool. Soc. | No. 1461a,| No. 2933, = # R.C.g.| Specimen. | B. Mus R.C.8 : ail in. in. in. in in Length of skull from tip of nasals to middle of occipi- ORE CFUM. ope cd ccnicvesecs 23 21°875 21 °925 21°25 22 125 Breadth across nasal bones| 5°25 6 375 _ 4°75 6°75 0 eal of 2nd upper molar Wiis tanddnetaces 2 1-775 17 1°6 Length of Ist upper molar seevas eubucaeese 1°75 15 1°375 1°35 a cacy of 4th upper pre- :- molar at base........... 1°6 1°25 1°255 1-225 The following is an account of the various features of the viscera that were observed in the Sumatran Rhinoceros. K 2 eS re *- Page 94. 132 THE VISCERAL ANATOMY Alimentary Canal.—The palate, which is covered with a smooth epithelium, is marked by conspicuous transverse angular ridges. There are eleven of these on each side; and they are not continuous anteriorly across the middle line, but the prominences of one side are carried on as the fossee on the other. The posterior ridges are con- tinuous from oue side to the other; and they, instead of being trans- verse, as are those in front, are arranged in the form of a V, the concavity of the V being directed forwards. On the soft palate, which is 43 inches long, these ridges disappear. The palate is narrow, as can be readily seen by an inspection of the skull; anteriorly its breadth is 2$ inches, and posteriorly 3} inches, gradually increasing from before backwards. ; In no part of the alimentary canal, except on the surface of the tongue, were there any papille visible to the naked eye. The cheeks form pads on either side, composed of areolar and mus- cular tissue, which project into the cavity of the mouth. These pads are shaped like prominent blunted triangles, with their apices directed Fig. 1. Tongue of C. sumatrensis (superior surface). , p, soft palate, embracing the root of the tongue pig, epiglottis. OF THE SUMATRAN RHINOCEROS. 133 forwards ; they are 33 inches deep behind, where they are lost on the fauces, and they are about 9 inches long. The epithelium covering them is nearly smooth, and is very thick. The tongue is elongate, and in shape much like that of the Rumi- nants, being thin from above downwards in front, and deep behind, with a somewhat sudden transition from one to the other. From the apex to the posterior of the circumvallate papille is 15 inches, and from the epiglottis to the same papille is 2? inches. In the middle of the anterior thin portion the breadth is 2¢ inches, and in the middle of the posterior moiety it is 44 inches. There are many circumyallate papille, 33 on one side and 26 on the other, forming two clusters, separated by a smooth median longi- tudinal line. Each cluster is triangular in shape; and the two acute- angled triangles they form lie side by side and have their apices directed backwards. The individual papille which go to form them are largest posteriorly, reaching a diameter of } inch; anteriorly they Page 95. get smaller, and cease by becoming more and more scattered. The rest of the tongue is covered uniformly with filiform papille, among which no fungiformes are to be seen. The soft palate runs downwards as well as backwards; and its posterior portion, as Professor Flower specially pointed out to me, so - closely embraces the base of the tongue that, except when in the act of swallowing, the epiglottis always projects quite into the posterior narial chamber, as in the horse and many other animals. The an- terior portion of the soft palate is 2 inch thick, and very glandular. A collection of glands of considerable size on each side of the fauces are the only representatives of the tonsils. The salivary glands present the usual characters. The parotid is much the largest. It weighs 1 Ib. 1 oz., and is of an irregular semi- lunar shape, the concavity embracing the superior portion of the angle of the jaw; it is mostly situated between the body of the masseter and the posterior insertion of the sterno-mastoid muscle. It lies almost entirely below the level of the zygoma, sending up a small portion into the interval between it and the external auditory meatus. Its duct, which is 14 inches long, commences at the inferior angle of the gland, and, as in the Ungulata generally, runs round the lower margin of insertion of the masseter muscle, and up along its anterior border till it pierces the buccinator, to terminate by a simple orifice in the well-marked fossa between the cheek-pad described above and the superior gum, in a line with the interval between the first and second upper true molar teeth. The submazillary gland weighs 24 oz., and is irregularly cubical in shape. It is situated just under the angle of the jaw, covered by the digastric muscle. The duct is 134 inches long; anteriorly it is Page 96. 134 THE VISCERAL ANATOMY closely bound to the inner surface of the sublingual gland; and it opens far forwards, close to the frenum of the tongue, on either side of it. The sublingual gland weighs 2 oz., and is composed of several small portions which open separately, almost in one straight line, about half an inch apart, below the sides of the tongue, and parallel with the ramus of the jaw. The whole gland is about 6 inches long and 1 inch deep. The esophagus is thick and muscular, not of large calibre; it has the mucous membrane but loosely connected with the muscular parietes, and arranged in bold longitudinal folds. The stomach is of a very different shape from that of the Indian Rhinoceros as figured and described by Professor Owen, and in most. respects resembles that of the horse. It forms a broad tube much bent upon itself, with the cardiac and pyloric orifices approximated, and a deep and narrow interval between them, in which the main vessels. and nerves run, and across which the peritoneum extends. There is no definite constriction between the cardiac and pyloric por- tions of the viscus; but there is a peculiar diverticulum from the outer portion of the cardiac extremity, of a subconical form, in which the base of the cone is the attached end. The whole organ is therefore somewhat globose, with the above-mentioned cardiac cecum projecting _ to the left side. Fig. 2. Stomach of C. sumatrensis (inferior or parietal surface). es, termination of cesophagus ; py, commencing duodenum, just beyond pylorus. OF THE SUMATRAN RHINOCEROS. 135 With regard to the size of the stomach, the greater curvature is 62 inches in length, and the lesser 6 inches; the greatest breadth from side to side, including the cardiac diverticulum, is 26 inches; the greatest depth is 18 inches; and the length of the diverticulum is 11 inches, whilst it is 5} inches in diameter. The diameter of the undis- tended cesophagus where it enters the stomach is 2 inches, and of the commencing duodenum 1} inch. There is not a trace of enlargement of the duodenum at its pyloric end, like the considerable dilatation in the Indian species. A large portion of the pyloric portion of the stomach is situated beyond, or to the right of, the pylorus itself; but it is only a direct continuation of the cavity of the viscus, and hardly forms a true cul-de-sac. The great omentum, which does not cover the intestines, is of considerable size; it contains no fat, and is not in any way attached to the colon, but runs up, behind the stomach, free to the vertebral column. The interior of the stomach presents a similar condition to that found in the Tapir and Horse, the mucous membrane being of entirely different characters in the cardiac and pyloric portions. That in the cardiac end, and in its diverticulum, is much plicated in all directions, and has a white opaque appearance; while the pyloric portion is covered with a thick and apparently smooth mucous membrane of the Page 97. - ordinary colour. The line of junction of these two portions is abrupt; Fig. 3. < Stomach of C. sumatrensis (inner surface). ; es, esophagus ; py, pylorus; ce, cardiac cul-de-sac. 136 THE VISCERAL ANATOMY and its position can be best understood from the accompanying draw- ing, in which it is seen that the corrugated white opaque epithelium only covers about one-fourth of the whole cavity—namely, the margins of the cesophagus for about an inch, and the diverticulum, from which it extends to the right, and backwards for a short distance. The walls of the stomach are nearly uniform in thickness, being a little more muscular at the cardiac extremity and along the lesser curvature than elsewhere. When the organ is fully distended the diverticulum be- comes less conspicuous, the direction of its superficial fibres being from its base to its apex. The pyloric muscular ring is strong and nearly an inch thick, projecting into the tube. The small intestine is 36 ft. long, and of a nearly uniform circum- ference of 6 inches, reaching 7 inches in the duodenum. For the first six inches after the pylorus the mucous membrane is smooth and simple, much like that in the pyloric portion of the stomach. The seventh and eighth inches present irregular folds, which immediately give place to a perfectly uniform series of thin, continuous (or nearly continuous), transverse foldings, just like the valvule conniventes of the human Fig. 4. Mucous membrane of the small intestine, natural size, showing the valvule conniyentes. OF THE SUMATRAN RHINOCEROS. 137 small intestine. There are nineteen of these folds in each six inches of the intestine ; and they continue unchanged to within half an inch of the Page 98. ileo-cecal valve, where they cease. Their great number (over 1,300), . extreme simplicity, and uniformity is very striking; they project nearly $ inch into the intestine. Many are continuous right round the tube; but where two approach one another, as is frequently the case, an intermediate one frequently ceases after having made nearly a complete circle. A few are to be seen extending for only about an inch ; but most are either that size or considerably longer. There are no traces of any triangular or cylindrical papille the whole length of the intestine. The bile and pancreatic ducts open on a papilla situated a foot from the pylorus, among the valvule conniventes, on the mesenteric border of the gut. This papilla is conical and rounded, projecting half an inch, with a single orifice at its apex. There is a second smaller orifice for a duct two inches further on, between two of the valvule and on one side of the main one. No Peyer’s patches could be found; and in their usual situation there was no irregularity of the valvule conniventes. The ileo-czcal valve does not project to any extent into the colon; Page 99. but where the small intestine ceases, on the border of the ileo-cecal orifice which is nearest the caput ceci, there are two closely approxi- mated globose, apparently glandular masses, about the size of Tangerine oranges, situated in the walls of the intestine. The colon presents features of great interest, and agrees in its con- volutions with the Indian Rhinoceros. When the abdomen is opened by a ventral longitudinal and transverse incision, the posterior por- tion, or the hypogastric region, is seen to be occupied entirely by a Page 100. large, apparently globose viscus, which is the ventral wall of the cecum: anteriorly to this, in the umbilical region, is seen a very capacious and sacculated tube, running nearly transversely and a little backwards as it tends to the left side; this is the posterior moiety of the enormous loop of the first part of the colon (ascending colon in man). Further forward, in the epigastric region, and some- what covered by the ribs, is seen another transverse, but less con- siderable, sacculated tube, which is the anterior moiety of the same loop. There is no omentum covering these viscera. Nothing more can be seen without moving these parts. When the intestines are removed from the abdomen, the following disposition of the viscera is observed. From the huge subglobose cecum, which is median in position, with its axis slightly obliquely , backwards and to the left, the colon is directed forwards and to the Page 101. | right; but it almost immediately gives rise to the very considerable colic loop, which is directed first transversely to the left, and con- io i 138 _ THE VISCERAL ANATOMY tinues on obliquely backwards, the anterior returning portion of which returns to the right hypochondriac region, where its mesentery is very incomplete, and it is firmly bound down to the adjacent parietes. The transverse colon, running from this point, is situated quite above the colic loop, and is also bound down at the left hypochondrium as at the right. The third part of the colon (the descending in man) is very sinuous in its course; it ends by a very simple igi flexure, and is continued on as the capacious rectum. The cecum is 3 feet long, and of nearly the same diameter; it is pyriform in shape, and much like that of the Tapir, the blind end being the narrower. It is traversed by three long longitudinal bands, between which it is folded in large sacculi. The colon springs from Fig. 5. Inferior view of the colon of C. sumatrensis. ce, cecum ; ¢./, colic loop; ¢, transverse colon, placed above the colic loop to show it more clearly. OF THE SUMATRAN RHINOCEROS. 139 the anterior end of the cecum, and immediately makes a short sigmoid curve to the right. Its interior is lined with a smooth, simple, irregu- larly plicated epithelium, the folds of which are quite removed when the organ is distended. From the ileo-cecal valve to the anus is 16 feet ; and in its broadest part the colon is 39 inches in circumference. It is peculiar that, as in the Horse and also in the Tapir, the tube is of very different dia- meters in its different parts, the bend of the colic loop being very narrow in comparison with its main parts. The proximal 7 of the colic loop is sacculated, and, at its middle, 13 inches in diameter; but at its bend, where it is situated in the left iliac fossa, it is much smaller, being only 6} inches across, and not the least sacculated. It continues thus uniform on its surface, and gradually dilating for about 24 feet till in the epigastric region it again becomes sacculated and very capacious, reaching a diameter of Fig. 6. SS . _ Superior view of the colon of C. sumatrensis. si, small intestine ; ce, cecum ; c.l, colic loop, with the tranverse colon between it and the excum. Page 103. 140 THE VISCERAL ANATOMY 164 inches. From this point it rapidly reduces in the transverse colon, remaining somewhat sacculated, with only one longitudinal band, which is at the mesenteric border, till at the sigmoid flexure the diameter is 64 inches. The colic loop is just 5 feet long. There are no regular folds of the mucous membrane of the large intestine, but many minor ones, which disappear when the tube is distended. The rectum is nearly 7 inches in diameter. This arrangement of the colon is different from that of the Horse in that the portion corresponding to the ascending colon is longer in the latter. In the Horse and Tapir the colic loop is formed from the transverse colon, in this Rhinoceros more from the right hypochon- driac angle of that viscus. In the direction of the cacum, namely backwards and to the left, the Rhinoceros agrees with the Tapir and differs entirely from the Horse. The liver is not large, considering the size of the sain. It weighs slightly over 15 lb., is Aattened, and has no gall-bladder. Adopting Professor Flower’s method of describing this organ, all the main divisions are indicated, though most of the fissures are not deep. The left lateral lobe is the largest, and is overlapped by the left central along its median border. On the anterior surface the fissure between the two extends upwards to the left lateral suspensory ligament, and therefore nearly through its whole surface ; posteriorly it only extends up to about two thirds the distance. The median suture, between the left and right central lobes, extends halfway up the organ anteriorly and not quite so far posteriorly, where it is stopped abruptly by a transverse bridge of hepatic tissue. The left central lobe is triangular, prismatic, and elongate, coming to a point below on a level with the general contour-line. One flat surface of this prism, the largest, is directed forwards; and the other two are wedged between the left lateral and right central lobes. The right central lobe is less differentiated from the right lateral than those just described are from one another, the fissure only extend- ing upwards a short distance; and it is itself cleft to nearly the same extent near the middle of its truncated inferior border. The right lateral, the lobe second in size, is suboval and simple, with the margin entire. Mesially it slightly overlaps the right central lobe at its inferior corner. The caudate is a very considerable lobe, shaped much like the left central, but larger; it is elongate, ovate, prismatic, and pointed at its free end. The largest side is directed forwards; and the external margin of the right lateral overlaps it considerably. It is 15} inches long, the whole liver, when lying on a flat slab, measuring 22 inches across, and 14 inches from above downwards. In no part does it measure more than 33 inches from before backwards. ene EE OF THE SUMATRAN RHINOCEROS. 141 Fig. 7. re Anterior or diaphragmatic surface of liver of C. sumatrensis. re, right central lobe ; ri, right lateral ; Ic, left central lobe; UJ, left lateral ; c, caudate lobe; sp, Spigelian lobe ; 4v, hepatic veins. Fig. 8. re Posterior or abdominal surface of liver of C. swmatrensis. re, right central lobe ; r/, right lateral; Je, left central lobe; J, left lateral ; ce, caudate lobe ; sp, Spigelian lobe. Page 102. Page 104. 142 VISCERAL ANATOMY OF THE SUMATRAN RHINOCEROS. The Spigelian lobe is most peculiar, mainly consisting of a thin strip of hepatic tissue, 8 inches long, uniformly ? inch wide, and {inch deep. At its attached end it becomes somewhat larger, and presents a free border superiorly for about an inch. There are three large hepatic veins, which spring just above and behind this lobe, on their way to the vena cava. The pancreas is irregular, not large nor concentrated. The spleen is very thin and flat; it forms an elongated oblong, rounded at one end and squared at the other. Its length is 25 inches, — and breadth 8 inches. It is slaty in colour, and weighs 2{ Ib. The kidneys are flattened and oval in form. One is 6 inches broad by 2 inches long. They are nearly equal in size, and together weigh 10 lb. The hilum is linear, and on the inferior surface, not at the margin. They are lobulated externally, but not so much as the Seals. The heart presents no peculiar features. The whole organ weighs 10 lb. when emptied of clots. The annulus ovalis is well marked, and forms a considerable fold over the fossa ovalis. The commencing aorta, which is 4 inches long and 34 inches across, divides into two nearly equal branches, one of which is continued on as the arch of the aorta, with a diameter of 1°85 inch; the other gives off the vessels to the head in the following manner. Immediately after the main divi- sion of the vessel into two parts, the innominate gives off the left subclavian, which, again, is much divided up. The innominate then, 3 inches above its origin, divides into the right subclavian and the common carotid trunk, which latter, after a simple course of 23 inches, divides into the right and left common carotids. This disposition is very much like that of the Llama as drawn by Professor Owen. The thickness of the ventricular septum is 14 inch. The lungs are extremely simple, coniform, and undivided, except at their apices, where, as in many animals, they send down small lobes which overlap the auricles of the heart. They are nearly equal in size, being 25 inches long by 15 deep and 5 broad. They weigh each 94 lb. (uncongested). The uterus is two-horned. The corpus uteri is 3} inches long by 24 inches broad; the cornua are 16} inches, by 2 inches broad; they are both very distinctly longitudinally plicated. The os uteri is much folded, and the orifice is quite small; from it to the orifice of the urethra is 12 inches. The vagina is lined with a squamous epithelium, and it presents a few transverse folds about 3 inches apart. Its cir-- cumference in the middle is 15 inches, at its orifice 9 inches. The urethra is 2 inches long, and admits two fingers. The length of the elongate fringed orifices of the Fallopian tubes is 5 inches. The clitoris and vulva are similar to those of the Indian species. THE BRAIN OF THE SUMATRAN RHINOCEROS. 143 23. ON THE BRAIN OF THE SUMATRAN RHINOCEROS Page 411. (CERATORHINUS SUMATRENSIS).* [Plate IV.] Ty a communication to this Society, published in its ‘‘ Proceedings” in 1873 (p. 92), I had the opportunity of describing the visceral anatomy of the Sumatran Rhinoceros (Ceratorhinus sumatrensis) from the first specimen received by the Society. A second individual of the . species, a female (as was the first), was deposited in the Gardens by Mr. C. Jamrach in July 1875, and was subsequently purchased. It unfortunately died on May 30th of this year, with symptoms of lung disease, a post-mortem examination demonstrating that both lungs were uniformly and throughout implicated. My friend Dr. James F. Goodhart, of Guy’s Hospital, late Pathological Registrar at the College of Surgeons, has kindly examined these organs, and reports to me that they “show a very extensive catarrhal pneumonia, degenerating in the centres of most of the patches. There is, in addition, some peribronchial inflammation, evidenced by a large growth of nuclei in the submucous and deeper tissues of the bronchi. The disease there- fore precisely corresponds with the caseous pneumonia to which man is subject.” The specimen is the one referred to by Mr. Sclater in his valuable and superbly illustrated memoir in the Society’s “‘ Transactions,” vol. ix. p- 651 (foot-note %). Feeling how important it is to obtain all possible information with reference to the species, and not haying removed the brain in the earlier specimen, I took the opportunity of doing so in the second, and on the present occasion place before the Society the drawings of the brain from different aspects (Plate 4, [LX X]), for verification of which I would refer the reader to the Museum of the College of Surgeons, _ where the original will be found preserved and mounted. The brain of the Indian Rhinoceros (Rhinoceros unicornis) is repre- sented in its different aspects, and in its internal detail, by Professor . Owen, in the “ Transactions” of this Society, vol. iy. pls. 19—22, and is described shortly on page 58 et seq. of the same volume. To this it is my desire that the figures here given should form a companion. By comparison it will be seen at a glance that the brain of Rhinoceros unicornis is slightly more simple than that of Ceratorhinus * “Transactions of the Zoological Society,” X. pp. 411-3, pl. LXX. Read, June 19, 1877. Page 412. 144 THE BRAIN OF THE SUMATRAN RHINOCEROS. sumatrensis, although the greater size of the former species would have favoured an opposite conclusion. So complicated and numerous are the convolutions that the general type-plan of their disposition is to a considerable extent disguised. They very closely resemble the same in the Equide, as might have been surmised. The whole brain, however, is broader, especially near the posterior portion of the cerebral hemispheres, where the breadth is considerably greater than further forward. The accompanying diagram will facilitate the description. It re- presents the disposition of the main conyolutions upon the superior aspect of one hemisphere, and exhibits the direction of the sulci which divide them. Two diagonal sulci cut up the posterior part of each lobe into three oblique gyri, which may be called the (1) external, (2) middle, and (3) internal gyrus. The middle and internal of these fuse together near the transverse line which joins the two rudimentary Sylvian fissures, anteriorly to which there is, in the Equide, no indi- cation of further primary longitudinal division. The external oblique gyrus continues, from this line, directly forwards, and independent. Upper view of left cerebral hemisphere of Ceratorhinus sumatrensis, showing general direction of sulci. In Ceratorhinus sumatrensis the internal oblique gyrus is triangular in shape, its inner boundary being the great longitudinal fissure of the hemispheres, into which it descends a short distance. In the Equide the inner boundary of this gyrus is more superficial, and can be seen as a straight longitudinal line, just external to the fissure itself, in the superior view of the brain. The whole gyrus is much broken up by minor foldings of its elements, especially in its median portion, its outer moiety consisting of a minor gyrus, whose general direction is THE BRAIN OF THE SUMATRAN RHINOCEROS. 145 a continuous oblique line, fairly regularly bent upon itself, first one way and then the reverse. The median oblique gyrus is divided into two nearly equal moieties by a fissure running parallel to its direction, each half being much - doubled upon itself. Anteriorly bridging minor convolutions blend it with the internal oblique gyrus, about one third distant from the ante- rior extremity of the hemisphere, in front of which the broad oblong cerebral surface is divided by a longitudinal suleus into two equal moieties, both convoluted. In the great breadth and division of this anterior portion the Rhinoceros under consideration differs from the Equide, and agrees with Rhinoceros wnicornis. The external oblique gyrus is much doubled on itself, and sepa- rated from the Sylvian fissure, which it surrounds, by minor convolu- tions, more strongly differentiated anteriorly. On the inner surface of the hemisphere the hippocampal gyrus is seen to be traversed by minor sulci and slight folds which ran parallel to its length, as in the Equide, the eaJloso-marginal sulcus following the anterior bending of the corpus callosum, and not, as in so many Artiodactyla (but not in the Equidew), becoming superficial ante- riorly. ‘ The fissure of Sylvius forms an open angle, at the bottom of which Page 413. are situated a number of small convolutions radiating from a point, which I take to be the island of Reil. The under surface of the brain exhibits the smooth surfaces of the middle lobes of the hemispheres and the smooth broad roots of the equally broad olfactory nerves, which are not lobate at their anterior extremities. The optic chiasma is short, the two optic nerves springing from its anterior surface quite close together. The pons Varolii is not large, the reverse being the case with the crura cerebri and the corpora albicantia. The lateral lobes of the cerebellum are small compared with the median portion, as is the case in the Ungulata generally. DESCRIPTION OF PLATE 4 (LXX), Brain of Ceratorhinus sumatrensis. a Fig. 1. Lateral view of right cerebral hemisphere. 2. Inferior view of left half of brain. 3. Superior view of left half of brain. 4. Internal view of right cerebral hemisphere. rer 146 SOME POINTS IN THE VISCERAL ANATOMY OF THE Page 24, ON THE DEATH OF A RHINOCEROS IN THE. SOCIETY’S GARDENS, AND ON SOME POINTS. IN ITS ANATOMY.* “ Mr. A. H. Garrod, in drawing attention to the death on Decem- ber 14th of the female Rhinoceros unicornis, which had lived in the Society’s Gardens for more than twenty-three years, remarked that the only pathological sign detected was the enlargement of the lymphatic glands at the base of the heart. Mr. Garrod’s observations on the visceral anatomy of this Rhinoceros were quite confirmatory of those of Professor Owen. In addition, he mentioned that there was a minute os cordis at the attached margin of one of the aortic valves, and that in the Perissodactyla this bone is not always absent, as by: some supposed, he having found a large one in a Sumatran Tapir. The remarkable difference between the arrangement of the mucous membrane of the small intestine in the Indian and Sumatran Rhino- cerotes (that of the former being produced into villi nearly an’ inch long through its whole length, whilst'in the latter’ these were repre- sented by valvule conniventes) was also illustrated from specimens in: spirit.” Page 707.25, ON SOME POINTS IN THE VISCERAL ANATOMY OF THE RHINOCEROS OF THE SUNDERBUNDS (RHINOCEROS SONDAICUS)+4 Our present knowledge of the visceral anatomy of the Rhinocerotide is confined to that of the two species Rhinoceros wnicornis and Cerato- rhinus sumatrensis. Professor Owen has given us, in the “ Transac- tions” of this Society (vol. iv. pp. 31 et seq.) an exhaustive account of the former of these animals; and-in the “ Proceedings” (1873, pp. 92 et seq.)~ it has been my endeavour to indicate most of the im- portant features in the latter, which, as Professor Flower has kindly pointed out to me, were briefly described by Sir E. Home in the * “ Proceedings of the Zoological Society,” 1874, p. 2. Read, Jan. 6, 1874. + “Proceedings of the Zoological Society,” 1877, pp. 707-11. Read, Nov. 6, 1877. t Supra, p. 130. RHINOCEROS OF THE SUNDERBUNDS. 147 * Philosophical Transactions”’ (1821, p. 271). On the present occa- sion I bring before the Society my notes on a young female of the Sondaic Rhinoceros (Rhinoceros sondaicus), which died in the mena- gerie of Mr. C. Jamrach, after having been in this country for a little more than half a year. It was only the skinned trunk which came into my possession. It is the nature of the mucous membrane of the small intestine which was certain to be of greatest interest; and this I am able to describe in detail. The individual under consideration measures, stuffed, 6 feet 2 inches from the tip of the nose to the base of the tail. The tail itself is a foot long, whilst the height of the animal at the shoulder is 3 feet. From the middle of the occipital crest, along the curve of the superior surface of the skull, to the tips of the nasal bones is 13} inches, the same measurement in adult animals being 22 inches. The single milk-incisor on each side of each jaw is still in place, as are all the milk-molars. The first true molar has not cut the gum; but its cap is seen within the bony alveolus. No traces of the other molars are visible. Mr. E. Gerrard lias kindly lent me the skull for examination. In Page 708. its base it exhibits the characteristic peculiarities of the species so clearly enunciated by Professor Flower,* the vomer being free behind - and developed into a tongue-shaped process ; the mesopterygoid fossa being expanded, and the free ends of the pterygoids everted at the same time that they are broad. No second combing-plate is present on the uncut first upper molar tooth. The animal is too young to be contrasted advantageously with Professor Peters’ drawing+ of Rhinoceros inermis, Lesson. I have, however, taken the opportunity of comparing that figure with the skulls of R. sondaicus in the College of Surgeons’ Museum, and fail to see that there are sufficient differences to justify specific differentia- tion. Professor Flower had previously done the same, and had arrived at a similar conclusion, as he found that even greater differences than those pointed out by Professor Peters are to be detected in individuals which are all undoubtedly of Indo-Malay origin. In skin-folding and surface-texture the Sunderbund and Javan Page 709. specimens agree exactly ; the young Sunderbund animal presenting a most striking uniformity in the size of the epidermic tuberculation, except in the gluteal region, where the boiler-bolt-shaped tubercles are somewhat larger than elsewhere. Along the back the scattered brown hairs, which spring from the yielding linear intertubercular surfaces, _ave also well developed. * “ Proceedings of the Zoological Society,” 1876, p. 447. + “Monatsb. der konigl. Akad. zu Berlin,” 1877, p. 68, pl. ii. L 2 148 SOME POINTS IN THE VISCERAL ANATOMY OF THE is 1. uf W Ly bic rial i us a Cie wx aS is ean ~ wy ww Mucous surface of duodenum of Rhinoceros sondaicus. : The following are the lengths of the alimentary viscera :— _ Small intestine, 26 feet 2 inches. Large intestine, 9 feet 10 inches. Cecum, 1 foot 3 inches. The stomach, in shape, is very much lke that of R. unicornis as figured by Professor Owen. Its cardiac surface is lined with the smooth white squamous epithelium found in all the Perissodactyla. This occupied about one third of the total gastric area, extending along most of the lesser curvature, the rest being covered with a Page 710. smooth and thick digestive coat. There is no trace of any cesophageal valve like that found in the Horse. The small intestine is somewhat larger in the duodenal region than elsewhere. Its first three inches are destitute of the flattened papille found elsewhere; but here, as all along the small intestines, minute villi are present everywhere. Three inches from the pylorus the papillee commence, and resemble those similarly situated in Rhinoceros unicornis,* except that they are not quite so long. They are repre- * Vide Professor Owen’s figure, “Transactions of the Zoological Society,” IV. pl. XII. fig. 1. LS RHINOCEROS OF THE SUNDERBUNDS. 149 sented in Fig. 1 [p. 148], where they are seen to consist of flattened, round-tipped processes of the mucous membrane, several of which are blended at their bases, in transverse lines. None are more than ‘3 of an inch in length, and most about “6 inch broad where they first become free. They give the impression of being incomplete valvule conniventes which have been cut and deeply jagged at their free edges. The opéning of the bile-duct is 7 inches from the pylorus, being a nipple-like tubular projection, nearly an inch long, among the papille. From the spot where they commence, all the way to the ileo-cxcal valve, these papille are found—those near the last-named situation differing from those in the duodenum in being more scattered and freer from one another, many in the ileum springing independently from the mucous membrane. Nowhere, however, are they otherwise Page 711. than flattened, broad, and blunt-tipped, none anywhere being circular and slender like those in the ileum of R. unicornis,* the existence of which I have had the opportunity of verifying. They never exceed Fig. 2. Mucous surface of ileum of Rhinoceros sondaicus. ‘3 of an inch in length. Numerous Peyer’s patches exist in the ileum, as may be inferred from Fig. 2, which is a representation of * Transactions of the Zoologieal Society,’ IV. pl. XII. fig. 3. 150 THE VISCERAL ANATOMY OF THE RHINOCEROS. a portion of the inner surface of the small intestine quite close to the ileo-ceecal valve. Such being the case, R. sondaicus differs from R. wnicornis in that the papille of the ileum are short, flat, and broad, instead of long, cylindrical and narrow, “ like tags of worsted” (Owen). . The cecum coli is a short blunt cone, with the diameter at its base as great as its length (1 foot 3 inches); and comparing the disposi- tion of the colic flexures and proportionate diameter, I found them identical with those of the Sumatran species as I have figured them.* Fig. 3. Liver of Rhinoceros sondaicus. Visceral surface. L.L. Left lateral. 2.C. Left central. 2#.C. Right central. #.L. Right sa lateral. C. Caudate. Sp. Spigelian lobe. The liver wants the gall-bladder, and differs but little from that of the Sumatran species. Fig. 3 is an outline-sketch of its abdominal surface, which, when compared with that of Ceratorhinus swmatrenis (“‘ Proceedings of the Zoological Society,” 1873, p. 102),¢ shows that the right central lobe is larger than the right lateral, instead of smaller. The Spigelian lobe is equally long and slender. The pancreas is of good size and fairly concentrated. The uterus is bicorn, each cornu measuring 8 inches, at the same time that the corpus uteri is 3 inches long. Each ovary is situated in a pocket of the peritoneum. * “ Proceedings of the Zoological Society,” 1873, pp. 99,100. (Supra, pp. 187, 138.) + Supra, p. 141. THE TNIA OF THE RHINOCEROS OF THE SUNDERBUNDS. 151 26. ON THE T4NIA OF THE RHINOCEROS OF THE Pace 78s. - SUNDERBUNDS (PLAGIOTENIA GIGANTEA, PETERS).* In 1856+ Dr. Wm. Peters described a tapeworm which he found in an African Rhinoceros from Mossambique, which he named Tenia gigantea. In 1870¢ Dr. Murie described the adult proglottides of a tape- worm passed by an Indian Rhinoceros (Rhinoceros wnicornis) living in the Society’s Gardens at the time, which he named Tenia magna ?. Tn 1871§ Dr. Peters communicated to the Society a Note on the results of a comparison of his specimens of Tenia gigantea with Dr. Murie’s description and figures of his Tenia magna?, showmg their identity, and suggesting the generic name Plagiotenia for the During this summer I have had the opportunity of eviscerating a half-grown female of Rhinoceros sondaicus, from the Sunderbunds, which had been a little more than six months in this country. In the - commencement of the colon I found three tapeworms with their heads (scoleces), together with several detached groups of proglot- tides, || these latter being quite undistinguishable’ from those figured by Dr. Maurie, in form as well as size. Dr. Peters has figured the scolex in his species, which is evidently in a powerfully contracted condition, to which one of my three specimens closely approaches. My other two specimens are not so, and, as a result, differ so much im appearance that I subjoin a figure of one of them. : Scolex of Plagiotenia gigantea, much enlarged ; superior and lateral view. * “Proceedings of the Zoological Society,” 1877, pp. 788, 9. Read, Nov. 20, 1877. t+ “ Monatsb. der Akad. der Wissensch. zu Berlin,” 1856, p. 469. ft “ Proceedings of the Zoological Society,” 1870, p. 608. § “ Proceedings of the Zoological Society,” 1871, p. 146. || In his account of his specimens Dr. Murie has most curiously mistaken the groups of proglottides (which he figures) for single segments. Page 789. Page 196. 152 THE ANATOMY OF THE BINTURONG. Of the specimen here figured the breadth (after being kept in aleohol) of the scolex, opposite the suckers, is 4 millimetres, whilst the depth, to the lower of the two more strongly marked transverse lines below the suckers (the proliferating area), is 3 millimetres. The breadth of the largest of the proglottides is 3:1 centimetres, their depth being 4°5 millimetres. One decimetre from the end of the scolex the proglottides are 1°42 centimetre in breadth. In one respect the scolex differs from that described by Dr. Peters, the rostellum or little conical elevation between the suckers being scarcely even indicated as such. This, however, seems hardly suffi- cient to justify specific separation. It is an interesting fact that three different species of Rhinoceros so separated in their distribution, should be troubled with the same tapeworm, which must therefore, unvarying, have followed the ancestral species in its different variations, now so easily distinguish- able. 27. NOTES ON THE ANATOMY OF THE BINTURONG (ARCTICTIS BINTURONG).* CrrTaAIN points in the anatomy of the soft parts of the Binturong, a knowledge of which is necessary to assist in substantiating the generalisations of Mr. H. N. Turnert and Professor Flowerf as re- gards the correct classification of the Carnivora, being as yet unde- termined, the recent death of a male specimen enables me to supply them. Dr. Cantor§ and Professor Owen|| have described the alimentary canal, noting some of the most important points; but neither has entered much into detail, and the generative organs in the male are not included in their descriptions. Alimentary Canal. With regard to the palate, there are ten transverse ridges extend- ing across its anterior part; they are not very strongly marked. The * “ Proceedings of the Zoological Society,” 1873, pp. 196-202. Read, Feb. 18, 1873. : t “Proceedings of the Zoological Society,” 1848, p. 63 et seq. t “ Proceedings of the Zoological Society,” 1869, p. 4 et seq. § “Journal of the Asiatic Society of Bengal,” 1846, p. 192. || “ Anatomy of Vertebrates,” 1868, III. p. 445. THE ANATOMY OF THE BINTURONG. 153 anterior five form continuous curves, convex forwards, the first being just behind the incisor teeth; the posterior five, starting from the sides forwards and inwards, turn suddenly backwards at right angles - to their former direction, and, meeting in the middle line, produce _ V-shaped patterns, with the concavities directed forwards; they are Page 197. also somewhat further apart than those in front, and have one or two rows of mammillated projections in the spaces thus left. The back part of the palate is not ridged ; and the uvula is represented by two slight projections, one on each side of the middle line, with a very shallow notch between them. The tongue is 3 inches long from the tip to the posterior of the circumvallate papille; its sides are nearly straight and parallel, con- verging slightly in front; at its base the breadth is 1 inch, and in front it decreases to 4 inch. The mucous membrane covering its lower surface and the floor of the mouth is smooth; and the superior edge of the frenum lingue is 14 inch from the tip, which latter is simply rounded. The mucous membrane of the superior surface, which is thickly set with papille, extends up to and slightly over the margins of the tongue in its anterior part, forming a thin-edged fringe all along the border. The anterior half of the superior surface is covered with easily visible, hispid, feline, retroverted papillz, par- ticularly large at the centre, diminishing in size laterally and for- wards, where, at the extreme margin, some fungiformes are mixed up with them. In the back part of the tongue the papille fungiformes are sparsely scattered among the diminished filiformes; and the papille circumvallate, nine in number and not equal in size, form the usual V, four on each side, with one median and posterior. Between these and the epiglottis the mucous membrane is soft and covered sparsely with thin cylindrical papilla, some of which reach 4 inch’in length; these are most uniform in diameter from end to end near the middle line, and towards the sides they become shorter and broader at their bases, till they blend with and become un- distinguishable from their filiformes. No ossified lytta could be found. - The parotid is slightly the largest of the salivary glands ; it is irre- gularly shaped and thin at its edges, where it is interpolated between the muscles. The submaxillary gland is egg-shaped, and about ? inch in average diameter; its duct runs far forwards on the floor of the mouth, opening within 3 inch of that of the opposite side, npon the symphysis of the jaw and closely bound to it, just behind the canine teeth and half an inch behind the incisors: The sublingual gland is elongate, and nearly as large as the submaxillary. . The stomach has a very peculiar shape, being elongated longitu- dinally, and consisting of a longitudinal cylindrical portion running =. = Page 198. 154 THE ANATOMY OF THE BINTURONG: backwards, and, after an abrupt bend, returning chestwards, the parallel tubes thus formed being closely approximate. By this arrangement, notwithstanding the considerable length of the lesser curvature, the cardiac and pyloric orifices are not far from one another; and they would be nearer were it not for the fact that the second or returning portion of the tube is a little shorter than the first. The cardiac portion of this stomach-tube has a diameter in the undistended organ of 1 inch, which gradually reduces to ¢ inch near the pylorus. A globose cardiac cul-de-sac throws the cesophageal opening quite to the right of that portion of the organ, and so brings it into contact with the commencing duodenum, which, before its first flexure, is a direct continuation forwards (chestwards) of the second or pyloric portion of the stomach. The greatest length of the un- distended organ, which is from the cardiac cul-de-sac to the middle Stomach of the Binturong. es, esophagus ; py, pylorus. of the bend above described, is 43 inches, and from the same bend to the pylorus is 2? inches; the cul-de-sac is 14 inch across. Dr. Cantor says, “The stomach is remarkably lengthened, cylin- drical, the parietes much thickened towards pylorus. CMisophagus enters close to fundus ventriculi, in consequence of which there is but a slight difference between the curvatures. Length along the greater curvature 1 foot 2 inches, along the smaller curvature 1 foot 1 inch.” Prof. Owen remarks of the stomach of the Lion that it ‘lies less THE ANATOMY OF THE BINTURONG. 155 transversely to the abdomen than in Man.” Ina Leopard Cat (Felis bengalensis) that I have lately dissected the shape of the stomach was almost.exactly the same as that of the Binturong; and it was simi- larly situated—namely, with its two moieties running longitudinally and not transversely. . _ The intestines are evidently much shorter in the specimen that I dissected than in those described by others, as may be seen from the following Table :-— Dr. Cantor’s Prof. Owen’s Present Specimen. Specimen. Specimen. ft. in ft. in. ft. in. Small intestine ..........-..... (ee 0) 7 se | 49 Large intestine i> 40 2 0 1 hl Da Padi yaks Ss acosackcases 0 OO 0 OF 0 Length of specimen, without tail 2 8 2 0 2 3} The bile-duct joins that from the pancreas for } inch before it Page 199. enters the intestine, which it does 24 inches from the pylorus, at the second bend of the duodenum. The intestines are thick, as in the Cats; and there are no permanent folds in any part of the mucous membrane of the alimentary tract. The cecum is situated, as in the - Felide generally, in about the centre of the abdomen, on the inferior surface of the diagonal portion of the intestine, which runs to the left hypochondriac region, and then, after, in Arctictis, dilating slightly at Fig. 2. Portion of the colon of the Binturong, showing the small cecum (c), and the dilatation at the angle of the colon, which is situated in the left hypochondrium. the angle, goes straight backwards to the rectum. It is very much like that in the Herons, being of considerably less diameter than the 156 THE ANATOMY OF THE BINTURONG. gut itself; the colon and small intestine are of nearly equal diameter and uniformly cylindrical. The omentum only covered the intestines to a small extent, not going more than half down the abdomen, The liver presents all the known lobes; and the left lateral, right central, and right lateral are large. The lateral fissures extend deeply into the organ. The right central lobe is considerably cut up; the fissure of the gall-bladder is deep; and a small supplementary lobule covers the fundus of that viscus on its abdominal surface. The left central lobe is much more conspicuous on the diaphragmatic than on the abdominal surface. The caudate lobe is larger than usual, and quadrangular, presenting the renal fossa well developed, and being perforated by the vena cava inferior. The Spigelian lobe is elongate- oval, pointed at its free end, and it does not reach as far as the left Fig. 3. “Liver of the Binturong. The various lobes are lettered as follows :—x1, left lateral; 1c, left central; Rc, right central; RL, right lateral; s, Spigelian; c, caudate; and GB is the gall- bladder. In order to facilitate comparison, the direction of the shading in the different lobes is varied, all parts of the same lobe being shaded in the same direction, whilst the lobes on each side are differently shaded ; the left lateral, however, so far over- laps the left central as to appear to be connected with the right central. THE ANATOMY OF THE BINTURONG. 157 margin of the liver. The general contour of the lobes is even, with a few slight irregularities now and then. The spleen is long and thin; it is 6} inches long, ? inch across, tapering and rounded at the ends. The kidneys are smooth and reniform. Each lung is divided into distinct lobes, the left into three, and the right into four, the extra one on the right side (the azygos) being Page 200. behind and nearly in the middle line. The urinary bladder in this specimen was very much distended, and ascended a considerable way into the abdomen as a narrow pyriform sac. The testes were situated in the scrotum, which projected backwards from the greatly developed mass of perineal glands. Of these last- mentioned glands Dr. Cantor remarks :—“ Between the anus and penis is situated a large pyriform gland, exceeding 2 inches in length, partially divided by a deep naked fossa, commencing from the latter organ. The gland secretes a light-brown oily fluid, of a peculiar intense, but not fetid or sickening odour.” The deep cleft above mentioned is longitudinal; and it is over its naked approximated sides that the orifices of the numerous simple, pyriform, yellowish, translucent glands open. Each separate gland is about 4 or ¢ inch - long, and 2 inch across at its broadest part, near its base. The two Page 201. lateral aggregated collections of these glands make up the oval or nearly circular mass in front of the testes; and the raphe of the perineum runs at the bottom of the cleft between them. The penis, in its non-erect condition, does not project more than } inch beyond them. The prostate is present, but only forms a small glandular mass round the sides and inferior portion of the urethra. It is situated { 334 inches from the base of the bladder, being simple, 4 inch broad, and 2 inch long. Cowper’s glands are situated 1} inch in front of it; they are oval, and each is ,5; inch broad and $ inch long. The testes measure 13 inch by 4 inch. There is no os penis. The glans penis is conical and pointed, 7 inch long, and presents round its base several small dark brown hard flattened papille, about 2, inch long. The vesicule seminales are absent. The anal glands are simple, globose, and thin-walled, about 4 inch in diameter; their orifices, one on each side, are extremely small. The brain presents the feline characters so clearly pointed out by Prof. Flower;* and, as in Felis, it differs from that of Viverra in — * “ Proceedings of the Zoological Society,” 1869, p. 478. i 5 a 158 THE ANATOMY OF THE BINTURONG. having the posterior and not the anterior of the limbs of the internal circumsylvian gyrus of greater breadth. Fig. 4. Brain of the Binturong. The Sylvian fissure tends to be vertical, but is directed somewhat backwards as well as upwards. It is surrounded by three gyri. The inner commences behind, near the lower border of the temporal lobe, and, after ascending as high as the top of the fissure, bends round it and descends on the frontal lobe to the supraorbital fissure, when it again doubles forwards to form the commencement of the middle gyrus. Its posterior limb is twice the breadth of the anterior, and is bisected by a vertical fissure which extends down as far as the hori- zontal temporal fissure. The middle gyrus is of uniform breadth throughout, and, commencing at the folding of the inner gyrus on the frontal lobe, goes round it and terminates at the lower border of the temporal lobe behind; there is no fold in it at its posterior superior angle. The third or outer gyrus is but slightly bent in its anterior Page 202. limb, which commences at the supraorbital fissure; it embraces the middle gyrus, and does not cease opposite its posterior superior angle, but descends about halfway down iis posterior limb to end by a point. The whole brain narrows in front; and the crucial sulcus is not at all strongly marked. The corpora albicantia are separated behind ; and the optic nerves in front of the chiasma run forwards close to- gether. The pituitary body is of fair size. THE ANATOMY OF THE BINTURONG. 159 28. NOTE ON THE ANATOMY OF THE BINTURONG Page 142. (ARCTICTIS BINTURONG).* Iy an earlier communication + I was able to confirm the observations of others, and to add fresh details, with reference to the anatomy of Arctictis binturong. Since that paper was published, two other speci- mens of the species have passed through my hands, the earlier of which differed in no respect from the one which I have previously described. The last, however, which died on January 4, 1878, pre- sented a peculiarity which I feel to be deserving of record. It was a _ female, apparently adult, having lived in the Gardens of the Society since October 19, 1875. The abnormal feature which it presented was the total absence of any trace of the colic cecum, which, as is shown in a drawing accompanying my former paper, is normally extremely small. The line of separation between the small and large intestines is well defined ; and there is no valvular constriction between the two tubes, as is the case in the Arctoidea generally. There is a large Peyer’s patch quite close to the termination of the small gut. The non-constancy of the presence of the diminutive cecum in Arctictis binturong, and its total absence in Nandinia binotata,t makes it evident that the existence of the cecum is a less important diag- nostic character than was inferred by earlier investigators. * “ Proceedings of the Zoological Society,” 1878, p. 142. Read, Feb. 5, 1878. + “Proceedings of the Zoological Society,” 1873, p. 196. (Opp.) t Vide Prof. Flower's “ Note,” “ Proceedings of the Zoological Society,” 1872, _ p. 683. Page 202. 160 THE CAUSE OF DEATH OF A BLACK-FACED KANGAROO. 29. ON THE CAUSE OF DEATH OF A BLACK-FACED KANGAROO (MACROPUS MELANOPS).* Tue cold weather of the first week of this month coming on rather suddenly, seems to have been the cause of the death of three animals in the Gardens, in all of which, on post-mortem examination, it was found that the lesion was the result of excessive and abnormal move- ment in the abdominal viscera. A Paradoxure died from intussuscep- tion of the small intestine, part going through the ileo-czcal valve into the colon; an Emu from prolapse of a considerable length of the ali- mentary canal; and the above-named Kangaroo from strangulation of a loop of small intestine by the tight twisting round it of the csecam— a most uncommon lesion, which proves that the possession of that ap- pendage has its disadvantages as far as the individual is concerned— just as in several human subjects death has been proved to have occurred from impaction of small bodies, like cherry-stones, in the appendix vermiformis. In the Kangaroo under consideration, on opening the abdomen the attention was immediately drawn to a large loop of strangulated small intestine, quite black from congestion, and partly covered with flakes of recent lymph, the result of the induced peritonitis, which was in- considerable. The length of gut involved was nearly two yards after it had been detached from the mesentery; but in the body of the animal it appeared considerably shorter, from being convoluted in the ordinary manner. The last foot or so of the small intestine was not included in the diseased loop, which consisted of the portion immedi- ately preceding it. The cecum was about a foot and a half long, and was situate in the right iliac region, from which it extended to the left superficially, and then again to the right behind the loop of intestine which it encircled, so that the caput ceeci could be seen, distended with grumous matter (as was the strangulated portion), to the right. With care, while the viscera were in situ, the little finger could be introduced into the ring thus artificially formed; and it was evident that the constriction was mostly produced by the mesenteric band which attaches the proximal portion of the. cecum to the small in- testine. There were no adhesions of importance. The viscera were removed en masse; and afterwards, without the least difficulty, the cecum was uncoiled, and the intestine was then left quite pervious. * “Proceedings of the Zoological Society,’ 1873, pp. 202-3. Read, Feb. 18, 1873. THE CAROTID ARTERIES OF BIRDS. 161 The mesenteric border of the cecum was nearly as black as the stran- Page 203. ' gulated part; but it was more normal in colour elsewhere. The ali- mentary canal was not at all over-distended with food; and the colon was nearly empty. Till the attack came on which caused its death, the animal was in excellent health. It was ill only forty hours. At first it lay out straight on its back for some hours; but during the last day of its life it was much doubled up, with its head between its legs. 30. ON THE CAROTID ARTERIES OF BIRDS.* Page 457. Between the years 1825 and 1830 three anatomists published the results of their independent observations respecting the number of and the variations in the carotid arteries of the different members of the class Aves. The first of these was Bauer,t who, in 1825, pointed out some of the most noteworthy peculiarities, which have been sub- sequently verified. Meckel,f in 1826, was enabled to demonstrate the existence of other marked variations; and his observations, extending over a considerable period, are incorporated in his “ Comparative Anatomy.” In 1829, C. L. Nitzsch selected the same subject for a disquisition before the University of Halle.§ Since that time scarcely any further additions have been made, and the subject has been almost entirely neglected. It is not easy to understand the reason of this; for it is generally acknowledged that what has been already done by the above-named authors is extremely valuable as an assistance to- wards a knowledge of the correct classification of birds, and yet they have left much for other workers in the same field. The opportunities afforded me by this Society, as their Prosector, and by many kind friends, who have supplied me with specimens, in spirit, of genera and species otherwise unobtainable, have enabled me'to collect together a sufficient number of facts, previously unrecorded, to make me feel justi- fied in presenting to this Society a fresh list, in which is recorded the arrangement of the carotids of the various birds examined by myself, at the same time that the previously known results of Bauer, Meckel, * “Proceedings of the Zoological Society,” 1873, pp. 457-72. Read, May 6, 1873. f Disquis. cirea nonnullarum Avium systema arteriosum (Berol. 1825). } “ Beitrag zur Geschichte des Gefiiss-Syst. der Vigel,” Meckel’s Archiy, 1826. § Observationes de Avium arteria carotide communi (Hale, 1829). M q; : : haa 7 é t - Page 458. 162 THE CAROTID ARTERIES OF BIRDS. and Nitzsch, and a few others, are incorporated in the general state- ments. In birds, the aorta, immediately after it has sprung from the heart, divides, as stated by Meckel, and contrary to the opinion of Cuvier, into two branches, the left innominate and the continuation of the main trunk, This latter again almost immediately divides into the right innominate and the descending aortic arch. Each innominate, after sending off pectoral and subclavian branches, continues to ascend a short way; and when near the superior aperture of the thorax it divides into the carotid, vertebral, and thyroid branches, except in those in which the carotid of one side is deficient. In what may be called the typical arrangement, the carotids, equal in size or nearly so, run up the front of the neck from the inner side of each thyroid Fig. 1.* Fig. 2. h. Fig. 1. Carotids at the base of the neck in aves bicarotidine normales. Fig. 2. Carotids at the base of the neck in aves levo-carotidine. gland, converging until they meet in the middle line, at which spot they enter the median intermuscular septum, and continue up to the head, on the front of the bodies of the remaining cervical vertebra, in the hypapophysial canal, covered by the lateral cervical muscular masses, and, where they are present, threading the bony arches. Birds with this arrangement are said to have two carotids, and may be termed aves bicarotidine normales (see Fig. 1). * In these diagrams, which represent the main arteries at the root of the neck, the following is the explanation of the abbreviations :—A, origin of the aorta at the heart; @, arch of the aorta; J.i, left innominate artery; 7.2, right innominate artery; 7.s, left subclavian, and 7.s, right subclavian artery; J.c. left carotid, and r.c, right carotid artery. THE CAROTID ARTERIES OF BIRDS. 163 A second group is peculiar in having the right carotid branch of the innominate undeveloped, when the left only traverses the hypapo- physial canal, being of large size; it bifurcates shortly before it reaches the head, thus producing a vessel on each side, to be dis- tributed in the same way as the terminations of the carotids in the previous group. Such birds are said to have a left carotid, and may be termed aves levo-carotidine (see Fig. 2). In a third arrangement, found only in certain Parrots (see Fig. 3), the right carotid artery runs in the hypapophysial canal, and the left at the side of the neck superficially along with the corresponding pneumogastric nerve and jugular vein. Birds with this arrangement Page 459. may be termed aves bicarotidine abnormales (see Fig. 3). Fig. 3. Fig. 4. A Fig. 3. Carotids at the base of the neck in aves bicarotidine abnormales. Fig. 4. Carotids at the base of the neck in aves conjuncto-carotidine. Fourthly, the two carotids, running apparently as usual, directly they meet, join and continue as a single trunk till near the head, where the single vessel bifurcates, as in birds with a left carotid only. These may be termed aves conjuncto-carotidine. In the common Bittern, where this condition obtains, the arteries (Fig. 4) are equal in size or very nearly so; but in the Flamingo (Fig. 5) the left is ex- tremely small, and has been on this account overlooked by previous observers, Meckel, Nitzsch, and Owen stating that there is only a Page 460. right carotid in Phanicopterus. I have had the opportunity of examin- ing two specimens of Pheenicopterus antiquorum and two of P. ruber ; and in all of them both carotids were present in the lower part of the neck, the right being much the larger and being joined by the left to form one trunk at the point in the neck where they first meet, as in Botaurus stellaris. Both vessels carried blood; but the calibre of the M 2 164 THE CAROTID ARTERIES OF BIRDS. left was extremely small, and that of the right was nearly the same as it would have been if it alone had been present. . ae Fig. 5. Fig. 6. Fig. 5. Carotids at the base of the neck in the genus Phenicopterus, as found by myself in all specimens. Fig. 6. Carotids at the base of the neck in Cacatua sulphurea, according to Meckel. From the list at the end of this paper it is shown that of 300 genera in which the arrangement of the carotids has been observed, in 193 of them both are present, in 107 the left only ; in one only are both equal when they join in the neck; in another they join, the left being the smaller; and in one other the right is the smaller under similar con- ditions; whilst perhaps one possesses the right only. So it may be generally stated that in birds either both carotids are present sepa- rate, or the left only exists. Several attempts have been made by different authors to account for these peculiarities. According to Bauer, the simplicity of the carotids (in other words, the presence of the left instead of two) is dependent on the size of the individual, the smaller species having the single trunk. Undoubtedly the great majority conform to this rule; but there are too many exceptions, as shown by Meckel, to make the generalization of much value, Rhea, Podiceps, Cacatua, Talegalla, and Menuwra possessing only the left. Meckel originally thought that a correlation existed between the length of the neck and the simplicity of the carotids; but when he found two carotids in Struthio, Dromeus, Cygnus, and Ardea he acknowledged that such was not the case. Prof. Owen remarks*, “ Birds as a rule are peculiar in sleeping with their long necks much bent or twisted; and this position might be expected to exercise some effect on the vessels subject thereto. Accordingly we find that the * “ Anatomy of Vertebrata,” vol. ii. p. 190. THE CAROTID ARTERIES OF BIRDS. 165 carotids are frequently of unequal size; in the Dabchick* the left is the largest; in an Emu I found it the smallest.” I may here remark that on several occasions I have watched the Flamingos sleepmg; and they do so, some with the neck bent one way and some the other, in a manner quite independent of the constant peculiarity in the arteries of their necks. All these explanations, therefore, fail to show why birds should have two or only one carotid artery ; and it is the last of them only that takes into consideration which carotid would be absent when there is any deficiency. If it were proved that all birds with a left carotid slept with their necks bent in one direction, the only explana- tion would be, that they did so because the arrangement of their cervical vessels would not allow of their doing otherwise, and con- sequently an argument in a cirele would be the only result. The ulti- mate cause is most probably as yet some way beyond our grasp; but I would offer the following as a step towards it. In birds possessing Page 461. two carotids those vessels, after they have once met, run close to- gether in the hypapophysial canal, but do not blend or anastomose in any way. In Botaurus stellaris, Cacatua sulphurea (according to Meckel, as shown in the diagram, Fig. 6 [p. 164]), and the genus Phenicopterus, - the carotids join to become one vessel at the spot where, in others, they come into contact, each proximal portion persisting. What I desire to show is, that on simple mechanical principles it is much more ' likely, when the two vessels do so blend, that the right should disap- pear, leaving the left solely to maintain the cerebral and cervical cir- culation; in other words, the assumption that there is a blending of the left with the right carotid in early life is sufficient to explain the absence of the right in birds thus affected. The diagram, Fig. 4, [p. 163] (which shows the distribution of the arteries at the base of the neck as they would appear immediately after the fusion of the caro- tids), will help to explain my meaning. The blood-current, almost immediately it has passed the aortic valve, divides into two, one going along the left innominate, and the other following the course of the aorta until it very shortly further divides into that traversing the right innominate, and that which continues on to the abdomen and posterior extremities. Such being the case, and the two carotids being of equal calibre, it is evident that, just as in Wheatstone’s Bridge the electric current is less intense in the bridge itself than in the branches, the current in the right carotid, which, in the case under consideration, connects the left carotid with the aorta distad of the point at which * In the Grebes (Podiceps), according to my observations, the right carotid is not found to be present at all—A. H. G. Page 462. 166 THE CAROTID ARTERIES OF BIRDS. the left innominate springs, is less than in the vessels it connects; consequently the current there tends to stagnate; but a tendency to stagnate in blood is a tendency to coagulation, as is seen in the proximal end of a ligatured arterial trunk; and the tendency to coagulation is a tendency to obliteration of the vessel in which the coagulation occurs ; consequently the right carotid must tend to dis- appear, which it does in nearly every case. Since this explanation occurred to me, I have not had the opportunity of examining any of the birds in which the right artery persists after it has fused with the left, to see if there is any peculiarity in their vascular arrangement which will account for its persistence. When the carotids do not blend there is evidently no reason why either should disappear; and when they do join, the presence of a large pectoral and subclavian branch from each innominate does not alter the problem; it only indicates that the obliteration must occur distad of it, as is the case. The following list includes all those species of birds in which I have had the opportunity of observing the disposition of the carotid arteries. They are arranged nearly according to the classification adopted in Mr. Sclater’s revised List of the Vertebrated Animals in the Gardens of this Society. PASSERES. All the Passeres examined possess the left carotid only. Species examined. OSCINEs. a. Oscines dentirostres. Turdus merula. grayt. Sylvia hippolais. Luseinia vera. Erithacus rubecula. Pratincola rubetra. Ruticilla phenicura, Myjiadestes obscwrus. Sialia wilsonit. Troglodytes parvulus. Mniotilta varia. Oinclus aquaticus. Motacilla flava. Hirundo rustica. Anthus pratensis. Parus major. Sitta ewropea. Lanius collurio. Sigmodus caniceps. Struthidea cinerea. Oriolus, sp. Artamus, sp. Graucalus mace. Dicrurus leucops. Muscicapa griseola, Ptilogonys cinereus. Ampelis garrulus. b. Oscines latirostres. Ohelidon urbica. THE CAROTID ARTERIES OF BIRDS. 167 c. Oscines tenuirostres. Nectarinia, sp. Prosthemadera nove-zealandie. Zosterops albogularis. Tropidorhynchus, sp. Diceumn, sp.° Diglossa baritula. Anthornis melanura. Cereba cyanea. d. Oscines conirostres. Tanagra cana. Donacicola castaneothoraz. Euphonia violacea. Oyanospiza ciris. Cissopis leveriana. Cardinalis virginianus. Estrelda melpoda. Coccothrastes vulgaris. Quelea occidentalis. Hedymeles ludoviciana. Buplectes capensis. Pyrrhula vulgaris. Ploceus manyar. Corythus enucleator. Hyphantornis castaneo-fuscus. Linaria cannabina. Padda oryzivora. Emberiza, sp. Munia maja. Alauda arvensis. Poéphila cincta. Melanocorypha calendra. e. Oscines cultrirostres. . . Icterus abeiller. Ptilonorhynchus holosericeus. : Molothrus bonariensis. Heteralocha gouldi. “ Cassicus persicus. Corvus coraz. Ageleeus ludovicianus. — frugilegus. Sturnus vulgaris. australis. Gracula religiosa. Garrulus glandarius. Cyanocorax cyanopogon. Strepera graculina. Page 463. Cissa speciosa. TRACHEOPHON2. Pitta, sp. Pitangus sulphuratus. Rupicola crocea. : Hylactes megapodius. Tipaugus cineraceus. Menura superba, Tyrannus satrapa. MACROCHIRES. TROCHILID2. The left carotid only is present in Patagona gigas. Chlorolanypis osberti. CYPSELIDZ. The left carotid only is present in the following species :— Cypselus apus. Cypselus alpinus. 168 THE CAROTID ARTERIES OF BIRDS. Cheetura vauai. Chetura caudacuta. —— spinicauda. Dendrochelidon coronata. But both carotids were found to be present in a specimen of Oypseloides fumigatus. CAPRIMULGIDA. Both carotids are present in these birds. Species examined. Caprimulqus europeus. Chordeiles texensis. STEATORNITHIDZ. Both carotids are present in Steatornis caripensis. PICI. Picipz. The left carotid only is present in the Woodpeckers. Species examined. . Picus major. Chloronerpes yucatanensis. minor. _ Melanerpes formicivorus. Picoides tridactylus. Mulleripicus fulwus. Tiga jawensis. Gecinus viridis. Leuconerpes candidus. Yune torquilla. COCCYGES. CorRAcIID2z. Both carotids are found in these birds. Species examined. Ooracias garrula. Eurystomus, sp. P 4. peat TROGONIDA. The left carotid only is present in these birds. Species ewamined. Trogon mexicanus. ’ Trogon puella. MEeRopIDz. The left carotid only is present in.these birds. Species examined. Merops apiaster. Merops ornatus. Momorip2. Both carotids are present in these birds. Species examined. Momotus lesson. Humomota superciliaris. THE CAROTID ARTERIES OF BIRDS. GaALBULIDZ. Both carotids are present in these birds. Galbula albirostris. Urogalba paradisea. ALCEDINIDZ. | Both carotids are present in these birds. Species examined. - Alcedo ispida. Ceryle amazona. Halcyon, sp. —— mazima. Dacelo gigantea. Cittura cyanotis. —— cervina. BUCEROTIDZ. Both carotids are present in these birds.* . Buceros rhinoceros. Buceros coronatus. —— plicatus. — atratus. b - * 2 > Urvurw2. The left carotid only is present in Upupa epops. MusopHaciD2. Both carotids are present in these birds. Musophaga violacea. Corythaiz albocristata. Schizorhis africana. CucuLipz. Both carotids are present in these birds. Speci ned Cuculus canorus. are Centropus senegalensis. Cacomantis sepulcralis. Guira piririgua. Chrysococcyz, sp. Pheenicophaes, sp. RAMPHASTID2. The left carotid only is developed in these birds. Ripon ae Ramphastos cuvieri. Ramphastos carinatus. 169 * PS., Sept. 4—In a specimen of Toccus melanoleucus just dissected I find the left carotid only present. Page 466. 170 THE CAROTID ARTERIES OF BIRDS. CAPITONIDA. The left carotid only is present in these birds. Species examined. Megalema asiatica. Indicator major. Barbatula duchaillu. PSITTACI. The Parrots are peculiar for the variation that occurs in their caro- tids, which show four different arrangements: first, there may be two in the normal position; secondly, the right may, as it usually does, traverse the hypapophysial canal, whilst the left, in a manner quite exceptional, runs superficially along the side of the neck in company with the left pneumogastric nerve and the left jugular vein; thirdly, the right may be very small, and blend with the much larger normally situated left (in Cacatua sulphurea, according to Meckel) ; and fourthly, the left may alone be developed, as in the Passeres. The first of these four conditions is only found in Old-World Parrots; and the last two are restricted to the Cacatuine. Species examined. In the following species the first plan prevails, the two carotids running normally :— Stringops habroptilus. Prioniturus, sp. Calopsitta nove-hollandice. Hos cardinalis. Eolophus (Cacatua) roseicapillus. indica. Euphema pulchella. Trichoglossus concinnus. splendida. Loriculus, sp. bourkit. Aprosmictus scapulatus. Melopsittacus undulatus. Paleornis alexandri. Agapornis roseicollis. The following species belong to the second division, the right carotid running normally, whilst the left runs up the side of the neck, together with the left pneumogastric nerve and jugular vein :— Ara macao. Nestor notabilis. Conurus cruentatus. hypopolius. wantholeemus. Brotogerys tiriacula. jendaya. ——— virescens. petit. tui. holochlorus. Pionus menstruus. Caica melanocephala. Chrysotis festiva. Psitiacus erithacus. ochrocephala. THE CAROTID ARTERIES OF BIRDS. 171 Chrysotis levaillantii. _ Platycercus eximius Psephotus hematogaster — ‘ Cyanorhamphus auriceps. Psitiacula passerina. — nove zealandie. Lathamus discolor. In the following species, forming the fourth section, the left carotid only is developed :— Cacatua galerita. Cacatua cristata. I have not yet had an opportunity of examining the third con- _ dition, #.e. that said to occur in Cacatua sulphurea. ; ACCIPITRES. Both carotids are present in all these birds. Species examined. CaTHARTID2. Archibuteo lagopus. Cathartes atratus. araaien SSnrsgg Gyparchus papa. aliaétus lla. Ea —— vocifer. VULTURIDZ. Aquila nevioides. = Neophron percnopterus. —— audaz. ; Gyps Sulcus. Spilornis cheela. Thrasaétus harpyia SERPENTARIIDZ. Falco peregrinus. Serpentarius reptilivorus. —— melanogenys. Hypotriorchis subbuteo Fatconipe. Tinnunculus alaudarius. Polyborus brasiliensis. Melieraz monogrammicus. Milous ictinus. Astur palumbarius. Buteo vulgaris. Circus cineraceus. Sreicipz. Both carotids are present in these birds. Species examined. Striz flammea. Bubo poensis. Otus vulgaris. fasciolatus. Syrnium aluco. Ketupa javanensis. nebulosum. Scops zorea. Bubo mazimus. Athene noctua. virginianus. —— passerina. ——- bengalensis. —— brama. 172 THE CAROTID ARTERIES OF BIRDS. Glaucidium, sp. Surnia funerea. Pulsatriz torquata. - Page 467. STEGANOPODES. Both carotids are present in these birds. Species examined. Fregata aquila. Phalacrocorax carbo. Sula bassana. Phaéthon, sp. HERODIONES. ARDEIDA. Both carotids are present in the following species :— Ardea cinerea. Ardea egretta. —-~ goliath. garzetta. . —— purpurea. candidissima. alba. Nycticorax ewropeus. But the two carotids join at the lower part of the neck directly they meet in Botaurus stellaris. CICONIIDE. Both carotids are present in these birds. Species examined. Ciconia nigra. Leptoptilus crumeniferus. alba. PLATALEIDZ. Both carotids are present in these birds. Species examined. Ibis rubra. Ibis nippon. melanocephala. Platalea leucorodia. strictipennis. PHENICOPTERIDA. The right carotid is much larger than the left, which joins it low down in the neck. Phenicopterus antiquorum. Pheenicopterus ruber. ANSERES. Both carotids are present in all these birds. Species examined. Anser segetum. Bernicla canadensis. THE CAROTID ARTERIES OF BIRDS. 173 Chloéphaga, sp. Aiz galericulata. Cygnus nigricollis. Mareca penelope. —— buccinator. Dafila spinicauda. coscoroba. Querquedula erecca. Dendrocygna autumnalis. Metopiana peposaca. —— viduata. Fuligula cristata. —— fulva. Mergus castor. Tadorna rutila. —— albellus. COLUMBZ. Page 468. Both carotids are present in all these birds. Species examined. CARPOPHAGIDE. Geopelia humeralis. _ Carpophaga globicera. Turtur senegalensis. —— enen. —— aldabranus. Lopholemus antarcticus. Metriopelia melanoptera. Ptilopus melanocephalus. Chamepelia talpacoti. —- marie. Leptoptila jamaicensis. Treron calva. Chalcopelia ehalcospilos. y —— puella. 7 CoLumBIp2. Tympanistria bicolor. Columba enas. Ocyphaps lophotes. livia. Chalcophaps chrysochlora. —— leucocephala. Phaps chalcoptera. — picazuro. Phlogenas cruentata. —— maculosa. Calenas nicobarica. —— vinacea. Didunculus strigirostris. Geopelia striata. Goura coronata. —- placida. victorie. ’ —— cuneaia, GALLIN A. Both carotids are present in all this Order, except in the Turnicide and Megapodide, in which the left only is developed. Species examined. With both carotids. PTEROCLIDZ. Tetrao urogallus. : Pterocles alchata. arenarius. PHASIANIDZ, Francolinus vulgaris. TETRAONIDZ. afer. Tetrao tetraz. —-— ponticerianus. Page 469. 174 Francolinus gularis. clappertont. Arboricola torqueola. Perdiz cinerea. Coturniz communis. Rollulus coronatus. Odontophorus dentatus. Ortyx virginianus. Hupsychortyx cristatus. Caccabis chukar. Phasianus colchicus. versicolor. reevesit, Thaumalea picta. —— anherstie. Euplocamus erythrophthalmus. —— vieilloti. THE CAROTID ARTERIES OF BIRDS. Euplocamus pyronotus. —— horsfieldit. albo-cristatus. Gallus bankiva. Ceriornis temminckii. Pavo nigripennis. muticus. Argus giganteus. Meleagris gallopavo. Numida meleagris. CRACIDA. Orax globicera. incommoda. Penelope cristata. Ortalis albiventris. With the left carotid only present. TURNICIDE. Hemipodius tachydromus. MEGAPODIDA. Talegalla lathamt. Megacephalon maleo. ALECTORIDES. Both carotids are present in all this order. Species examined. OrIpDz. RAuuipz. TToubara macqueent. Gdienemus grallarius. bistriatus. Rallus aquaticus. Aramides cayennensis. Porzana americana. carolinensis. Orex pratensis. Ocydromus sylvestris. Porphyrio madagascariensis. CARIAMIDZ. Cariama cristata. Chunga burmeistert. melanotus. GRUIDE. Gallinula chloropus. Grus antigone. PARRIDA. EURYPYGIDZ. Parra africana, Rhinochetus jubatus. THE CAROTID ARTERIES OF BIRDS. 175 GRALLA. ; ‘Both carotids are present in these birds. Species examined. CHARADRIIDZ. Totanus calidris. Vanellus cristatus. _ —— solitarius. Charadrius pluvialis. Gambetta flavipes. — hiaticula. Machetes pugnaz. Hematopus niger. Scolopaz rusticula. Strepsilas interpres. Gallinago scolopacina. —— gallinula. ScoLopacip2. Tringa canutus. Numenius arquatus. cinclus. —— pheopus. Glareola, sp. Timosa lapponica. GAVL&. Both carotids are present in all these birds. Species examined. s LaRiDz. : Lestris antarcticus. Larus glaucus. Larus argentatus. Sterna hirundo. PROCELLARIIDZ. Page 470. Both carotids are present in all these birds. Species examined. Thalassidroma pelagica. 4strelata lessont. — fregata. Prion vitatta. —— bulweri. PYGOPODES. Both carotids are present in the following species :— CoLYMBID2. ALcID2. Colymbus glacialis. Alea torda. Uria troile. The left carotid only is present in the following species :-— CoLyMBiD2. ALcIDe. Podiceps cristatus. . ; Arctica alle. —— minor. Page 471. 176 THE CAROTID ARTERIES OF BIRDS. IMPENNES. Both carotids are present in these birds.’ Species ewamined. Spheniscus demersus. Aptenodytes pennantit. humboldtii. ; CRY PTURI. Both carotids are present in these birds. Species examined. Rhynchotus rufescens. Crypturus sallei. STRUTHIONES. Both carotids are present in Struthio camelus, The left carotid alone is present in Rhea americana. Both carotids are present in Oasuarius bennettit. Dromeus nove-hollandie. bicarunculatus. The left carotid only is present* in Apteryx mantelli. Apteryz owenii. In reviewing the above facts, for the purpose of forming an esti- mate as to the significance in classification of the arrangement of the carotid arteries in birds, the following conclusions may be drawn, if the Psittaci be omitted from consideration, which, from the great peculiarities they present among themselves, it will be better to do at present :— 1st. In many cases the uniformity of the carotids is of ordinal im- portance, all Passeres possessing the left carotid only, whilst both are present in all Columbex, Anseres, Gralle, and Accipitres. 2nd. As a family character, the distribution of the carotids is very suggestive, in some cases presenting a well-marked difference between * In his “ Anatomy of Vertebrates” (vol. ii. p. 199), Prof. Owen remarks :— “In the Apteryx I found that the left carotid alone passed to the usual place in the neck, and divided at the third cervical vertebra, to supply the head in the usual way.” But in the original memoir on the bird (“Transactions of the Zoological Society,” vol. ii. p. 278) the same author has stated that both are present, which is not the case in the specimens examined by me. eee THE CAROTID ARTERIES OF BIRDS. 177 families otherwise closely allied. For example, the Megapodide, together with the Cracidx, as Professor Huxley has so clearly shown,* from a well-established suborder of the Gallinz, and osteologically it is not easy to separate them; but in the Cracidz both carotids are present, whilst in the Megapodide the left only is found. The Phcenicopteride also present a condition peculiar to themselves. In the somewhat ill-defined group, the Coccyges, the carotids give rise to family characters of value. The Bucerotidze and Ramphastide differ in the latter possessing only a left carotid, whilst the former have both present ; and the affinities of the Upupidz may be considered nearer to the Ramphastide, on account of their agreeing with them in this point. The Apterygide, as well as the Turnicide and Podicipide, are also well distinguished from their allies by their single carotids. 3rd. That Struthio and Rhea must be more than generically distant from one another is indicated by many characters ; and the difference in their carotids favours their being placed in separate subfamilies ; no such difference tends to divide up the Casuariide in a similar manner. 4th. Respecting genera, there are none in which the peculiarities of the carotids are not constant in them; but there are some which are separated from others by a difference in the arrangement of these vessels. Cypseloides fumigatus, a Swift, apparently not at all peculiar otherwise, undoubtedly possessed, in the only specimen I have had the opportunity of dissecting, two carotids. That this was an indi- vidual peculiarity is extremely improbable, as no similar case has been recorded in any other genus; consequently this genus (or species as it may be) differs from all its allies, which only possess the left carotid. _ A similar case, resting on similar evidence (a single. specimen) is that of Arctica alle, which differs from Alea and Uria, with which its affinities are very close, in having the left carotid only, instead of both. Botaurus, amongst the Ardeide. has also an arrangement pecu- liar to itself. As previously remarked, the Psittaci present greater differences among themselves respecting the disposition of the arteries of their necks than all the other orders of birds taken together, one con- dition being peculiar to them, and the other conditions being all represented amongst them. Without entering into further details regarding these birds it is impossible to make any generalizations of Page 472. importance ; and I will leave the subject for a special paper on the order. It is not until the different conditions of the carotid vessels are * “Proceedings of the Zoological Society,” 1868,"p. 298. N Page 526 Page 527. 178 “SOME POINTS IN THE ANATOMY OF STEATORNIS. taken in connexion with the pterylosis, as well as the anatomy of the viscera and muscles, that a correct idea can be formed as to their true value in the classification of birds. The work of the illustrious Nitzsch assists much in this direction; and it is to be hoped that as facts become more numerous, ornithologists will realize that a correct arrangement will not be arrived at until anatomy is more thoroughly studied. In conclusion, I have to present my best thanks to Mr. Sclater for the kind way in which he has on all occasions throughout this inquiry assisted me with suggestions and advice—also to Professor Flower, Mr. O. Salvin, Mr. Sharpe, and Mr. Howard Saunders, for their so willingly putting at my disposal specimens in spirit of species which I should not otherwise have had the opportunity of examining. 31. ON SOME POINTS IN THE ANATOMY OF STEATORNIS.* TurovuaH the kindness of Prof. Flower, I have had the opportunity of examining two specimens of Steatornis caripensis preserved in spirit, as well as the skeleton of another; and Mr. Sclater has also kindly given me a skin to assist in the study of the pterylosis, and a nestling, which I have dissected. Many points in the osteology of this bird, as well as the description of the larynx, are to be found ina paper by Johannes Miillert ; and further details are given in the works of L’Herminiert, Sclater$, and Murie||._ The following notes relate almost entirely to the pterylosis and the anatomy of the soft parts, the skull being only described so far as to make it comparable with those in Prof. Huxley’s paper on the classification of Birds]. Pterylosis (fig. 1, p. 180).—AII the top of the head is covered with a scattered feathering, which is very much the strongest between the eyes. There is no tendency to the formation of longitudinal bands in * “Proceedings of the Zoological Society,” 1873, pp. 526-33. Read, June 3, 1873. + “ Miiller’s Arch. f. Anat.” 1842, p. 1-11, and elsewhere. t “Nouv. Arch, du Mus.” tom. iii. 1834, p. 321, and elsewhere. § “Proceedings of the Zoologieal Society,” 1866, p. 126. || “ Ibis,” 3rd ser. vol. iii. No. 9, p. 81. { “ Proceedings of the Zoological Society,”’ 1867. SOME POINTS IN THE ANATOMY OF STEATORNIS. 179 this region, like those in the Caprimulgidew. Above each eye there are two rows of closely set very stiff feathers, ranning parallel to one another and to the border of the upper eyelid, forming a double eye- brow. The upper of these is slightly the stronger; it is situated ¢ of an inch above the lower one, with a bare space intervening. The stiff feathers of which it is composed are slightly more than } of an inch long and are directed outwards. The lower eyebrow is } of an inch _ above the margin of the lid, which has no eyelashes and is bare: it does not extend quite so far forward or backward as the one above it; and its component feathers are not quite so long. The external auditory orifice is nearly circular and 3 of an inch in diameter; there is no operculum. It is surrounded by a single row of feathers, much like those of the eyebrows; they are all directed backwards, the anterior being slightly the longer and acting as a protection to the entrance of _ the ear. Several (about a dozen on each side) stiff simple vibrisse, many more than 14 inch long, spring from the side of the upper beak, and run directly forwards, partially covering the apertures of the nostrils. The dorsal tract, where it commences, is narrowed on account of there being a bare space above each ear; but when it reaches the upper part of the neck it broadens, and continues down the back of _ the neck as a not strong tract, which becomes narrower and stronger as it descends, till at a short distance above the tops of the shoulder- blades it is very strong indeed. It continues on in this condition, and bifurcates between the scapule to form a well-developed fork, with long branches, which become considerably weakened near their extremities. Between the lower ends of this fork the continuation of the dorsal tract commences, not connected with it at all, but quite free, as an upward-turned weak arrow-head, situated in the middle line. The axis or shaft of this arrow-headed tract, as it descends, becomes narrower and stronger till it ceases abruptly at the base of the long infundibuliform nude oil-gland, which closely resembles that of the Owls. In the upper part of the loins, above the arrow-head, at a short distance on either side of, and parallel with, the mid-dorsal tract, is a single row formed by four strong feathers, which are dis- tinctly separated from the rest. All over the loins, behind the Page 529. acetabula, there is a weak feathering which blends with the lumbar tracts. These last are consequently not very distinctly defined, and consist mainly of weakly feathered tracts, running from the knee obliquely downwards and backwards, leaving the tibize almost bare, with the exception of a few semiplumes which are scattered below the front of the knee. ; Between the rami of the jaws the large triangular surface is naked at the sides and weakly feathered along the middle line up to the nN 2 Page 528. ~ 180 SOME POINTS IN THE ANATOMY OF STEATORNIS. La) g : E Ss = s Pterylosis of Steatornis. symphysis (as in the Owls), where there are a few vibrisse, directed forwards, From this submaxillary feathered portion the inferior neck- tract springs; and behind the angle of the jaw a weak branch is sent up, on each side, to join the dorsal tract and head-covering behind the ears. A little lower down the inferior tract becomes more defined, SOME POINTS IN THE ANATOMY OF STEATORNIS. 181 though not strong; it continues simple as it descends, being of the same breadth as the lateral neck-spaces. Just above the upper or scapular extremities of the furcula it ceases in the middle line, leaving a bare interclavicular space; but it develops a branch on either side, which expands over the chest to form the pectoral tracts. The pectoral tract of each side is double, the inner of its divisions being the continuation of the main tract, which descends, narrow and strong, close to the carina sterni in its upper part, but further sepa- rated below, leaving over the epigastric region of the abdomen a con- siderable median space, which lower down is again reduced by their convergence to the anus, just in front of which they terminate. Each outer pectoral branch of the inferior tract is weak and very diffused, covering the sides of the body, leaving a narrow space between it and the main stem, except at the points just in front of the scapular ends of the furcnla, from which they spring, and below the inferior margin of the sternum, where they again blend, and continue down side by side, after their contact, nearly to the anus, the outer branch being the weaker and less defined. There is a weak hypopteral tract continued from the outer margin of the external pectoral branch. The under wing-surface is feathered along the forearm in several rows. The margin of the patagium is ~ thickly set with short strong plumes. The humeral tract is strong and separated by a narrow space from the well-covered upper wing- surface. ‘ There is no aftershaft to the feathers. There are ten primary remiges, and twelve secondary, of which the ten distal resemble each other, and the two at the elbow are reduced in size. The upper wing-coverts do not extend more than or quite so much as halfway down the secondary remiges. There are ten rectrices. The above described pterylosis clearly indicates that in the arrange- ment of its feathers Steatornis more closely resembles the Strigide than the Caprimulgide, though it differs considerably from both. It resembles the Strigide and differs from the Caprimulgide in having no aftershaft to the contour feathers, in not having the occipital tract divided up into narrow longitudinal rows, in having spaces on each side of the submaxillary tract, in haying the pectoral portion of the Page 530. inferior tract in two parts, of which the inner approaches the carina a sterni above and separates from it as it descends, in having the upper ; wing-surface uniformly feathered, and in having a large infundibuli- form oil-gland. In none of the Caprimulgide does the inferior tract continue simple down the neck, whilst in Striz flammea as in Steatornis it does not bifurcate till in the region of the furcula. But Steatornis resembles the Caprimulgide and differs from the Strigide in having —_——_ Page 531. 182 SOME POINTS IN THE ANATOMY OF STEATORNIS. ten rectrices. It differs from both, however, in that the inferior por- tion of the dorsal tract does not unite:at all with the scapular fork of the superior portion, in having the outer branch of the pectoral tract diffused and descending far over the abdomen, and in the general tendency to scattering of the feathers. | In the skull the lachrymal bones are not developed as they are in the Strigide and Caprimulgide. The palate is strongly desmogna- thous, as in the Falconidew, and much more so than in the Strigide, which are almost schizognathous. The palatine bones also meet across a Fig. 2. b Skull of Steatornis. a, base; b, superior surface. the middle line for } of an inch, in a manner which is quite peculiar, and can be best understood by a reference to the drawing, each bone being apparently folded on itself behind the point of junction with its fellow, and articulating with the basisphenoid rostrum, as well as anchylosing with the vomer by its inflected and upward-turned margin; each develops a very short slender anteriorly directed process close to the vomer, which projects forwards on each side of it near its middle. The vomer itself is a quarter of an inch long, slender and quite blended with the palatines; its anterior pointed extremity advances as far for- wards as the posterior border of the median palatine symphysis men- tioned above. The posterior external angles of the palatines, so large in Caprimulgus and Podargus, are not developed. The basipterygoid facets are large. In the eye the sclerotic ossifications are not consider- able, as in the Owls, being not at all unusually developed. SOME POINTS IN THE ANATOMY OF STEATORNIS. 183 In the atlas the cup for articulation with the occipital condyle is incomplete behind; and the odontoid process of the axis is situated near its posterior margin. In this conformation, the classificational importance of which was first pointed out by Mr. Parker, Steatornis agrees with the Strigide and the Caprimulgide, but not with the ‘Cypselide, in the one or two cases which I have had the opportunity of observing. The well-known peculiarities of the sternum do not seem to point definitely in any special direction; and in the other bones I have not observed any demonstrable tendencies. Digestive organs.—The tongue is thin, smooth, and triangular ; it is ¢ inch broad at -its base, and 3 of an inch long; the posterior angles are prolonged backwards for ¢ of an inch as angular processes with small papille on them; the posterior border is simple. The esophagus is capacious and uniformly cylindrical, with longitudinal plications in ‘its mucous membrane. The proventriculus is zonary and well deve- loped, the largest of its component glands, which are slightly racemose, being 3 of an inch long. The stomach forms a thin-walled, globose, capacious gizzard, with its mucous membrane, as usually, longitudi- nally plicated. The intestines are 22 inches long, capacious through- out, and especially so near the pyloric portion; the biliary and ~ pancreatic ducts open into it 24 inches from the pylorus, at the bend of the duodenal loop. The two intestinal ceca are 14 and 12 inch long, slender, and a little broader at the cecal than at the open ends; they are situated 2 inches from the cloaca. The trachea is a little more capacious above than below. As in many birds, the separate rings of which it is composed are not so deep in the middle line as they are laterally ; and as in each ring the upper and lower margins of one side in one ring, and of the other side of the next above and below, are slightly everted, whilst those of the other half are inverted to the same extent, when the rings are superimposed they produce the appearance seen in the accompanying drawing, as if each ring were narrow on one side and broad on the other. The syringe (fig. 3, p. 184), as has been described by others, is extremely peculiar, because it is formed in each bronchial tube, instead of at the bifurcation of the trachea. The trachea bifurcates at its lower end much in the same way that it does in Mammalia; and each bronchus continues down towards the lungs as a cylindrical or slightly flat- tened tube, composed of simple and entire rings of cartilage. In a specimen that I once saw, there were fourteen of these rings on each side; but in the one before me, which is figured here, the bronchi are not equal in length, the left bronchus containing thirteen and the Page 532. right ten complete rings above the commencement of the syrinx. Each semisyrinx, as it may be termed, is formed on the same principle 184 SOME POINTS IN THE ANATOMY OF STEATORNIS. as that of the combined organ in most of the non-singing birds. Taking for description that of the left side in the specimen figured, it — is there found that the thirteenth bronchial ring is complete, though Front view of the syrinx of Steatornis. considerably flattened from side to side; the fourteenth is not com- plete in the middle of its inner surface, it is a little longer from before backwards than the one above, and not so long as the one following it. The fifteenth is only a half ring, its inner portion being deficient; it is slightly convex upwards, and articulates, both at its anterior and pos- terior ends, with the fourteenth incomplete ring and the sixteenth half ring. The sixteenth half ring is coneave upwards, and so forms an oval figure in combination with the one above, which is filled with a thin membrane, to form part of the outer wall of the bronchus. There is a membrane also between the ends of these and the succeed- ing half rings, which completes the tube of the bronchus internally. The half rings which follow the sixteenth reduce in size, and are con- siderably smaller before they reach the lung. The lateral muscle of the trachea extends down the outer side of each bronchus, to be Page 533. attached to the middle of the first fully developed half ring. The depressor muscles of the trachea are independent of these. SOME POINTS IN THE ANATOMY OF STEATORNIS. 185 Steatornis has two carotid arteries, as have both the Strigide and . Caprimulgide. With regard to the myology of this bird, the only muscles which will be considered are those which have been found to have some bearing on the systematic position of birds generally. In the thigh, the ambiens (Sundevall)—the slender muscle which _ in many birds runs from the innominate bone, just above the aceta- bulum, along the inner side of the thigh to the knee, which it crosses obliquely in the fibrous capsule below the patella, and then blends with the flexor perforatus digitoram—is absent, as it is in the Strigide and Caprimulgide. The semitendinosus, the outer of the two muscles which form the lower fold of the thigh (the semimembranosus being the inner), and which runs from the region of the lower end of the innominate bone to the tibia, is present, as in the Caprimulgide, it being quite absent in all the Strigide. As in the Caprimulgide also, this muscle receives A Fig. 4. B Muscles at the outer side ide of the elbow ; A, of right wing of Caprimulgus europaeus ; B, of left wing of Steatornis. tpb, tensor patagii brevis; eer, extensor carpi radialis; 6, biceps; d, deltoid ; #, triceps ; &, humerus. an accessory head from the lower end of the femur, which helps te send a partial insertion of the muscle down the leg. The femoro-caudal (which runs as @ narrow muscular ribbon from the middle of the linea aspera of the femur to the coccyx, covered by the semitendinosus and crossed superficially by the sciatic artery and nerve) is quite absent ; it is well developed in the Caprimulgid, small in the Strigide, and absent in very few birds. In the upper limb the second pectoral (subclavius of Rolleston) is not large, extending about halfway down the sternum, as it does in Page 534. 186 SOME POINTS IN THE ANATOMY OF STEATORNIS. the Strigides, whilst in the Caprimulgide it is more developed, reach- ing the lower margin of that bone. The tensor patagti brevis—a muscle very constant in its insertion in the different families of birds, which arises mainly from: the superior extremity of the furcula on each side, and is inserted, after running in the patagial fold parallel to the humerus, into the outside of the fore- arm, near the elbow—in Steatornis agrees entirely with that of many of the Strigide, and differs slightly from that of the Caprimulgide, as may be seen from the accompanying drawings (fig. 4, p. 185), where, in the former, the main tendon becomes attached to the extensor carpi radialis longus directly, whilst in the latter it joins a second more superficial tendinous slip which runs back to the outside of the elbow, much as in the Passeres. By placing the above-mentioned facts in a tabular form, the com- parison between Steatornis and its allies will be more clearly seen. Steatornis. Strigide. - Caprimulgide. Number of carotids....... “2 2 2 Ambiens muscle..........| Absent........ | Absent.........| Absent. Semitendinosus muscle....| Present, with ac-| Absent.. ..| Present, with ac- cessory head... cessory head. Femoro-caudal-muscle ....| Absent......... Present Goal Present. Pectoralis secundus muscle. | Short .......... Short . . | Long. The semitendinosus is a muscle very constantly present in birds, being absent only in the Owls, Eagles, true Vultures, Humming-birds, and Swifts; consequently its presence in Steatornis is in favour of its being related to the Caprimulgide rather than to the Strigide. In endeavouring, from the facts recorded above, to form a correct notion as to the exact systematic position of Steatornis, the difficulties in the way are considerable. That it forms a family by itself there is little or no doubt, as it presents pterylographical and internal pecu- liarities found in no other birds. By a process of exclusion, an approximate idea of its position may be formed. The Strigide, Caprimulgide, Coraciide, Momotide, Galbulidw, and Steatornithide all agree in possessing the following characters—two carotids, well- developed cxca, a nude oil-gland, and no ambiens muscle. Among these, the Strigide differ from all the rest and resemble the Eagles, in having no semitendinosus; and the Steatornithide are equally pecu- liar in haying no femoro-caudal muscle. In its pterylosis, as shown above, Steatornis resembles the Strigide much more than any of the allied families, except that there are only ten rectrices. I have not dissected Podargus; but it agrees so closely with Caprimulgus in its pterylosis, according to Nitzsch, that it most probably must be included ON CERTAIN MUSCLES OF BIRDS. 187 in the same family. As Mr. Sclater has shown, Podargus has no oil- gland, that organ being very small in the Caprimulgide generally, but large in Steatornis. If the absence of the ambiens muscle in the Strigide has the sig- nificance which I put on it, and is sufficient justification, in conjunc- Page 535. tion with other differences, for the entire separation of this family from the other Accipitres, then the above mentioned group of families seems a natural one;* but if the Strigide are intimately related to the Falconide and Vulturide, it is so difficult to believe that the Coraciidee and their allies are related to the Falconide, that the entire separation of the Strigide from the Caprimulgide seems essential, in which case the position of Steaternis becomes more doubtful. 32. ON CERTAIN MUSCLES OF THE THIGH OF BIRDS, Page 626. AND ON THEIR VALUE IN CLASSIFICATION. (PL. ¥ : Part I+ In their works on the general anatomy of the animal kingdom Meckel and Cuvier have devoted special chapters to the myology of birds. The dissections on which their observations were based were evidently undertaken more with the desire to determine the relations borne by the muscles of birds to those of Mammalia and- Reptiles, than with the object of studying the variations in the arrangement of the muscles _ in the class itself. Nitzsch, Reid, Owen, Milne-Edwards, Coues, Selenka, and others have published their dissections of certain birds, as the Vulture, Penguin, Apteryx, Eagle, and Loon; and most of these are, from their accuracy and clearness, valuable additions to zoological knowledge. Sundevall seems to be the only ornithologist who has employed the variations that he has observed to be constant in different birds towards the furtherance of classification; and my results, on the D sence: discussed by him, in most cases correspond with his. The great opportunities afforded me by this Society for the study of a great many species of birds in the flesh, have reduced the diffi- * Prof. Newton has for.some time believed the Steatornithide and Caprimulgide . to be distinct families (¢f. Zool. Rec. vi. p. 67). + Part IL. “ Proceedings of the Zoological Society,’ 1873, pp. 626-44. Read, June 17, 1873. 188 ON CERTAIN MUSCLES OF BIRDS. culties connected with the dissection of any special soft part to a minimum; and in the present communication the results are recorded of my dissection of the region of the thigh of the birds which have passed through my hands. In this region there are six muscles, or well-defined portions of muscles, which may or may not be present; and my chief object has been to note their presence or absence, which in some cases is far from easy, as modifications may occur which disguise the true connexions of a muscle, and thus lead to misinterpretation. As a common Fowl happens to possess all these six muscles well developed and easily demonstrable, I will commence with a short Page 627. account of their condition and relations in it, from which a correct idea may be easily formed of their situation in other birds with the help of the accompanying details. In commencing the dissection of the leg of a Fowl in order most easily to observe the arrangement of the muscles about to be described, Fig. 1. Outer view of right thigh of Gallus bankiva, partially dissected. s, sartorius; v e, vastus externus; 40 and 07, biceps, origin and insertion; Z/, tensor fascie; fc, femoro-caudal; a/c, accessory femoro-caudal; st, semi- tendinosus; ast, accessory semitendinosus; sm, semimembranosus; Ad, ad- ductor; P, pubis; R, rectrices. ON CERTAIN MUSCLES OF BIRDS. 189 the body, from which most of the feathers have been removed, should be laid on its side, and a section made in the skin, in a line parallel to and just over the femur, along its whole length; from the ex- tremities of this line the sections should be continued at right angles to it, extending upwards and downwards from the end over the head of the femur, and along the outer side of the leg from that at the knee. The skin must be then dissected downwards as a flap off the muscular adductor mass below, and some way upwards above the level of the femur. After this has been done the following muscles will be found without difficulty :-— Tensor fascie.—This is the superficial muscle of the outside of the Page 62” thigh, covering the femur. It is flat and triangular in shape, and arises as a membranous expansion which covers the gluteus ii, from the lower two-thirds of the posterior border of the iliac fossa in which that muscle is situated, and from the fibrous septum which separates that muscle from the gluteus iii. Further down it has origin also from the whole length of the ridge which separates the postacetabular area from the external lateral surface of the ischium, and which may be termed the postacetabular ridge, as well as from the posterior border of the ischium, as far forwards as its junction with the pubis, being here slightly overlapped by the semitendinosus. The fibres - converge towards the knee; and the deep portion of the muscle blends in its course with the vastus externus, together with which it continues forward to become part of the broad thin tendon which covers the knee and is inserted in the front of the tibia-head, the patella being situated in it, together with the long, slender, and flat tendon of the ambiens muscle, which is situated below it, running obliquely from inside and above, outwards and downwards. In many birds, as the Falconide and Psittaci, this muscle does not extend below the level of the femur, but ends inferiorly by blending with the vastus externus; and consequently, where such is the case, it evidently cannot, as it does otherwise, cover any of the flexors of the leg. In the Bucerotide it is entirely absent. Whether this postacetabular portion of the tensor fascis is present or absent has some bearing on classification, as in the different families it is a very constant feature. Biceps cruris——The anterior portion of this muscle may be seen in the Fowl’s leg before the tensor fascie has been removed, just below it, near its insertion. This muscle is mostly covered by the tensor fascie, arising from the upper three-fourths of the postacetabular ridge, just in front of the origin of that muscle. Its fibres converge to form a round tendon, which in the outer side of the popliteal region is bent sharply downwards by passing through a tendinous sling which arises from the lower end of the femur, to be inserted on a Page 629. 199 ON CERTAIN MUSCLES OF BIRDS. prominence on the outer side of the fibula, about halfway down the leg. In the loop above this tendon, and consequently quite out of the way of compressing forces, one of the nerves to the leg and fout is con- tinued. In one or two birds, as Phaéthon, the biceps does not pass through any loop, but is inserted directly by a broad flat tendon into the upper part of the fibula. Semitendinosus.—This flat ribbon muscle runs amen parallel with the lower fibres of the biceps, just below it. Its origin is mostly from the tip of the transverse process of the first free coccygeal vertebra, and from the fibrous membrane between it and the inferior border of the ilium. Near its origin it, being superficial, curves over the posterior inferior angle of the ilium, and covers the inferior fibres of origin of the tensor fascie, running upwards and forwards towards the inner side of the head of the tibia, and so getting covered an- teriorly by the inserted end of the biceps. A rhomboidal sheet of muscle, arising from the anterior end of the linea aspera, descends to form an accessory head to this muscle, joining it anteriorly, on its outer side, by an oblique tendinous raphe, which continues down the back of the leg superficially. A small part of the main muscle, the inner, goes straight forward to end at the inner side of the upper extremity of the tibia-by a flat tendon; but most of it joins the acces- sorius to be continued down the leg. Some birds, as the Eagles and Owls, have no semitendinosus at all; some, as the Anserine birds and Penguins, have no accessory semitendinosus, in which case all the fibres go straight to the tibia-head ; whilst in most the above-described condition maintains. Semimembranosus.—This ribbon muscle runs parallel to, deep of, and next to the semitendinosus. It arises from the outer border of the anterior margin of the ischium for about a third of an inch, at the place where it is in contact with the pubis, the origin extending down to the lower end of the slight prominence at the point where the ischium slightly overlaps the pubis. It is inserted along with the tibial end of the semitendinosus into the inner side of the head of the tibia by a broad flat thin tendon. This muscle is very constant in birds: in the Grebes it is extremely thin, and may sometimes be absent, as stated by Sundevall; but I have seen it in some fresh specimens of Podiceps minor, though but very slightly developed. Ambiens.—This peculiar, small, but very long muscle is triangular or fusiform in shape. It arises from the tip of the short anteriorly directed spine which is situated just above the anterior border of the acetabulum, and runs along the inner side of the thigh to the inner side of the knee, where it is covered by the sartorius, which is above it in the former part of its course. Its thin tendon then crosses the knee, running in the substance of the fascial extensor tendon, just ON CERTAIN MUSCLES OF BIRDS. 191 in front of the patella to the outer side, where it joins the fibres of origin of the flexor perforatus digitorum. Femoro-caudal.—This long ribbon muscle is covered superficially by the tensor fascis and biceps above, as well as by the semitendi- nosus lower down. The sciatic artery and nerve cross it superficially at right angles close to its insertion as they course from the sciatic foramen, parallel to the femur, to the popliteal region. The femoral vein separates this muscle from the adductor muscles at their inser- tions, except in Dacelo, where it crosses the femoro-caudal superficially.* It arises from the (anterior) transverse processes of the two last coccygeal vertebrx, and is inserted into the linea aspera of the femur, at about one-third its length from the trochanter. An accessory head, arising from the upper thrée-fourths of the postacetabular ridge, and from the ridge which forms the lower margin of the origin of the obturator externus, joins the tendon of insertion of this muscle, and is also partly inserted into the linea aspera, between it and the head of the femur. It is thin, muscular, and broad, covering the obturator externus superficially, and is par- tially intersected by a fibrous sheet where it crosses its anterior border. The sciatic artery and nerve cross it superficially ; and the nerve to the semimembranosus is deep of it, whilst that to the semi- . tendinosus is superficial in some cases; the biceps completely covers it. Of the above-described muscles, five of them (the ambiens, the femoro-caudal, the accessory femoro-caudal, the semitendinosus, and Page 630. the accessory semitendinosus) vary; any one or more than one may be absent in different birds; and in my dissections my object has always been to record the conditions existing in the specimen under examination. The constancy of the peculiarities in the different indi- viduals of each species, in the species of each genus, and very ~ generally in the genera of each family, makes it evident to any one working at the subject, that much respecting the affinities of the different families of birds is to be learnt from the study of their myology, in connexion with the peculiarities of their other soft parts; and that these features will, in the long run, lead to a more correct classification than one based on the skeleton alone, becomes : almost equally certain. The variations in the five above-mentioned muscles form the subject of this communication, and the subjoined list contains the results arrived at by myself. A few of the facts now recorded will be found mentioned in the works of Meckel, Sundevall, and others. Re- * In Centropus phasianus the main artery of the leg is also the femoral, and not the sciatic, as in other birds ; it therefore runs with the femoral vein in that bird. Page 631. 192 ON CERTAIN MUSCLES OF BIRDS. ference to them is quite unnecessary, as they can easily be found in the works of those authors. The Passeres possess the femoro-caudal, the semitendinosus, and the postacetabular portion of the tensor fascise; the accessory semi- tendinosus is present in all except Dicrurus (which has only ten rectrices) ; the ambiens and the accessory femoro-caudal are absent.* The Pict possess the femoro-caudal, the semitendinosus, and the postacetabular portion of the tensor fascie; the ambiens and the ac- cessory femoro-caudal are absent; and they may be divided into two subfamilies, according to whether the accessory semitendinosus is present or absent. The accessory semitendinosus is absent in Picus major. Picoides tridactylus. minor. The accessory semitendinosus is present in Geeinus viridis. Chloronerpes yucatanensis. Leuconerpes candidus. Mulleripicus fulvus. Melanerpes formicivorus. _Hypowanthus rivolit. Yunz torquilla agrees in all these points with Gecinus viridis. The Steatornithide possess the semitendinosus, and a very narrow accessory semitendinosus. The ambiens, the femoro-caudal, the acces- sory femoro-caudal, and the postacetabular portion of the tensor fasciee are absent. Species examined. Steatornis caripensis. The Caprimulgide possess the femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascie; the ambiens and the accessory femoro-caudal are absent. Species examined. Caprimulgus europeeus. Chordeiles texensis. The Trogonide possess the femoro-caudal and the semitendinosus ; the ambiens, the accessory femoro-caudal, the accessory semitendino- sus, and the postacetabular portion of the tensor fasciz are absent. Species examined. Trogon mexicanus. Trogon puella. * In my paper on the carotid arteries of birds a long list is given of the Passeres in which the carotid vessels were examined ; in all these the muscles of the thigh . were dissected also. In a specimen of Pomatostomus temporalis there was an acces- sory femoro-caudal on the right side; on the left there was not a trace of it. ON CERTAIN MUSCLES OF BIRDS. 193 The Meropide possess the femoro-caudal, the semitendinosus, and the accessory semitendinosus; the ambiens, the accessory femoro- caudal, and the postacetabular portion of the tensor fasciz are absent. Species examined. Merops apiaster. Merops ornatus. The Cypselide and the Trochilide agree in possessing the femoro- caudal, at the same time that the ambiens, the accessory femoro- caudal, the semitendinosns, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz are absent. Species examined. Cypseloides fumigatus. Chetura spinicauda. Cypselus apus. Dendrochelidon coronata. —— alpinus. Chlorolampis osbertt. Chetura vauxi. Patagona gigas. The Coraciide possess the femoro-candal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascize; the ambiens and the accessory femoro-caudal are absent. Species examined. : Coracias garrula. Eurystomus, sp. The Momotide possess the femoro-caudal, the semitendinosus, and the accessory semitendinosus; the ambiens, the accessory femoro- caudal, and the postacetabular portion of the tensor fasciz are absent. Species examined. Momotus lessoni. Eumomota superciliaris. — equatorialis. The Galbulide possess the femoro-caudal and the semitendinosus ; the ambiens, the accessory femoro-caudal, and the postacetabular portion of the tensor fasciz are absent. Tn Galbula clbirostris the accessory semitendinosus is present; but it is absent in Urogalba paradisea. The Alcedinide possess the femoro-candal and the semitendinosus ; the ambiens, the accessory femoro-caudal, the accessory semitendino- sus, and the postacetabular portion of the tensor facie are absent. Species examined. Alcedo ispida. Dacelo cervina. Ceryle amazona. gigantea. Haleyon, sp. Cittura cyanotis. Page 632. 194 ON CERTAIN MUSCLES OF BIRDS. The Bucerotide possess the femoro-caudal, the semitendinosus, and the accessory semitendinosus; the ambiens, the accessory femoro-caudal, and the tensor fascis are absent. Outer view of right thigh of Buceros coronatus, dissected. b, biceps; s#, semitendinosus; ast, accessory semitendinosus; sm, semimembra- nosus; fc, femoro-caudal; ae, obturator externus; @, adductors; gl, 1, 11, and III, glutei; s, sartorius; s#, sciatic nerve; sa, sciatic artery; p, pubis. Species examined. Buceros rhinoceros. Buceros coronatus. plicatus. atratus. ——- bicornis. Toccus melanoleucus. The Upupide possess the femoro-caudal, the semitendinosus, and the accessory semitendinosus ; the ambiens, the accessory femoro-caudal, and the postacetabular portion of the tensor fascise are absent. Page 633. Species examined. Upupa epops. The Musophagide possess the ambiens, the femoro-caudal, and the accessory femoro-caudal, the semitendinosus, the accessory semiten- dinosus, and the postacetabular portion of the tensor fascie. . ON CERTAIN MUSCLES OF BIRDS. 195 Species examined. Musophaga violacea. ee albo-cristata. Schizorhis africana. The Cuculide possess the ambiens, the femoro-caudal, the semiten- dinosus and the accessory semitendinosus, and the postacetabular por- tion of the tensor fasciz; they may be divided into two subfamilies, according to whether the accessory femoro-caudal is present or absent. The accessory femoro-caudal is present in Centropus senegalensis. Guira piririgua. phasianus. Phenicophaes, sp. The accessory femoro-caudal is absent in Cuculus canorus. Cacomantis sepuleralis. Chrysococcy2, sp. The Ramphastide possess the femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascie; the ambiens and the accessory femoro-caudal are absent. Species examined. Ramphastos carinatus. Ramphastos cuvieri. -—— ariel. The Capitonide possess the femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciee; the ambiens and the accessory femoro-caudal are absent. Species examined. Megalema asiatica. Indicator major. Barbatula duchaillua. The Psittaci possess the femoro-caudal well developed, the semi- tendinosus, and the accessory semitendinosus; the accessory femoro- caudal and postacetabular portion of the tensor fasciz are absent. The ambiens may be present normally; it may be differentiated in the thigh, but fail to cross the knee, being lost on the fascia over it, or it may be absent. The ambiens is present in the following species :-— Psittacus erithacus. Ara macao. * Conurus zantholemus. chloroptera. jendaya. Caica melanocephala. N petat. Nestor notabilis. a — holochlorus. hypopolius. The ambiens is present, but does not cross the knee in eS Stringops habroptilus. 02 Page 634. 196 ON CERTAIN MUSCLES OF BIRDS. The ambiens is absent in :— Pyrrhura (Conurus) eruentata. Oalopsitta nove-hollandice. Brotogerys tiriacula. Cacatua galerita. virescens. * cristata. tovi. roseicapilla. twi. Calyptorhynchus banksit. Pionus menstruus. Eos cardinalis. Chrysotis festiva. indica. ochrocephala. Tanygnathus muellert. levaillantiz. Prioniturus, sp. Psephotus hematogaster. Platycercus eximius. Trichoglossus concinnus. Lathamus discolor. pallidiceps. Aprosmictus scapulatus. Psittacula passerina. Loriculus asiaticus. Agapornis roseicollis. chrysonotus. ~ Melopsittacus undulatus. Cyanorhamphus auriceps. Euphema splendida. nove-zealandic. —— pulchella. Paleornis torquata. bourkit. Accipitres.—The Cathartide possess the ambiens, the semitendino- sus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciee ; the accessory femoro-caudal is absent, and the femoro- caudal is present, though small, in Cathartes atratus ; it is absent in Gyparchus papa. The Serpentariide possess the ambiens, the accessory femoro- caudal, the semitendinosus and the accessory. semitendinosus; the femoro-eaudal and the postacetabular portion of the tensor fascis are absent. Species examined. Serpentarius reptilivorus. The Vulturide and Falconide agree in possessing the ambiens and the femoro-caudal; the accessory femoro-caudal, the semitendinosus, ‘the accessory semitendinosus, and the postacetabular portion of the tensor fascie are absent in all. Species examined. Milvus ictinus. Buteo vulgaris. Archibuteo lagopus. Helotarsus ecaudatus. Neophron percnopterus. Gyps fulwus. Vultur auricularis. Polyborus brasiliensis. ~ ON CERTAIN MUSCLES OF BIRDS. 197 Haliaétus vocifer. Falco melanogenys. — albicilla. lanarius. Aquila nevioides. Hypotriorchis subbuteo. —— audaz. Tinnunculus alaudarius. Spilornis cheela. Melieraz monogrammicus. Thrasaétus harpyia. Astur palumbarius. Falco peregrinus. Circus cineraceus. Page 635. Outer view of left thigh of Neophron percnopterus, partially dissected. 8, sartorius ; g, quadratus femoris; gl, glutei; 5 o and 5i, biceps, origin and inser- tion; fc, femoro-caudal; sm, semimembranosus; oe, obturator externus; ev, caudal vertebre ; P, pubis; BR, rectrices. The Strigide possess the femoro-caudal, which is always small ; the ambiens, the accessory femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascie are absent. Species examined. Striz flammea. Syrnium nebulosum. Otus vulgaris. Bubo mazimus. brachyotus. virginianus. Syrnium aluco. — bengalensis. Page 636. Page 637. 198 ON CERTAIN MUSCLES OF BIRDS. Bubo capensis. Athene passerina. poénsis. brama. fasciolatus. Pholeoptyna cwnicularia. Ketupa javanica. Glaucidium, sp. Scops zorca. Pulsatria torquata. Athene noctua. Surnia funerea. The Steganopodes must be considered in separate families. I have not dissected Pelecanus. Fregata aquila possesses the ambiens and the femoro-caudal, the latter being very slender; the accessory femoro-caudal, the semiten- dinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz are absent, Phaethon possesses the femoro-caudal (small), the semitendinosus (strong), and the accessory semitendinosus; the ambiens, the accessory femoro-caudal, and the postacetabular portion of the tensor fascie are absent. In this bird the biceps cruris is inserted into the fibula-head directly, without passing through a loop. ‘The family Phalacrocoracide possess the ambiens,* the femoro- caudal, and the semitendinosus; the accessory femoro-caudal, the accessory semitendinosus, and the postacetabular portion of the tensor fascie are absent. There is a peculiarity about the obturator externus of these birds, which is perplexing at first sight. In Sula this muscle, instead of, as usual, being inserted into the outside of the head: of the femur, has its attachment further forward, to the outer edge of the linea aspera, or the bony surface corresponding to it, midway between the head of the bone and the attachment of the femoro-caudal muscle. In Phalacrocoraz it is situated still further forward, being in contact by its anterior border with the posterior margin of the femoro-caudal, parallel to it in direction, and otherwise in situation exactly like an accessory femoro-caudal. However, that it is the obturator externus and not the accessory femoro-caudal is certain, from the facts that the nerve to the semimembranosus is superficial to it, that the whole upper part of its outer surface is covered with a tendinous layer, that none of its anterior fibres blend with the posterior margin of the upper end of the tendon of the femoro-caudal, and that the presence of this muscle is constant in birds. The accessory femoro-caudal is, on the contrary, superficial to the nerve to the semimembranosus; it is not tendinous externally up to its insertion; some of its fibres blend with the tendon of the femoro-caudal just before it joins the femur; and its presence is uncertain. In Sula the ambiens has a tendinous link at the outer * Meckel did not find the ambiens in the Cormorant; it is peculiar in that it runs through the substance of the large triangular patella, in a bon} canal. ON CERTAIN MUSCLES OF BIRDS. 199 Outer view of right thigh of Phalacrocorazr carbo, dissected. tf, tensor facie; gl Iv, gluteus quartus; oe, obturator externus; fc, femoro- caudal; Ad, adductor; 4, biceps; st, semitendinosus; sm, semimembranosus ; Pb, pubis. side of the knee, which runs upwards from it to the outer side of the anterior fibrous expansion over the joint. . Species examined. Sula bassana. Phalacrocoraz lugubris. Phalacrocoraz carbo. The Ardeide possess the femoro-caudal—though in some, as Ardea goliath, it is extremely narrow—the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascize which is very slightly developed; the ambiens and the accessory femoro-caudal are absent. Species examined. Ardea cinerea. Ardea egretta. goliath. garzetta. —— purpurea. Nycticoraz europeus. alba. Botaurus stellaris. Page 638. Page 639. 200 ON CERTAIN MUSCLES OF BIRDS. The Ciconiide possess the ambiens, the semitendinosus, and the accessory semitendinosus ; the accessory femoro-caudal is absent. The femoro-caudal is very small in Oiconia and absent in Leptoptilus ; and the postacetabular portion of the tensor fascie is the same. Species examined. Ciconia alba. Ciconia maguari. nigra. Leptoptilus crumeniferus. The Plataleide possess the ambiens, the femoro-caudal, the accessory femoro-caudal, the semitendinosus, the accessory semiten- dinosus, and the postacetabular portion of the tensor fascise, which is small. Species examined. Platalea leucorodia. Ibis melanocephala. ajaja. nippon. Ibis rubra. —— spinicollis. strictipennis. The Phenicopteride possess the ambiens, the accessory femoro- caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascie; the femoro-caudal is absent. Species examined. Phenicopterus antiquorun. Pheenicopterus ruber. The Anseres possess the ambiens, the femoro-caudal, the acces- sory femoro-caudal (very large), the semitendinosus, and the post- acetabular portion of the tensor fascie; the accessory semitendinosus is absent. Species examined. Anser segetum. Tadorna rutila. Bernicla canadensis. Aix galericulata. Chloéphaga, sp. Mareca penelope. Cygnus nigricollis, Dafila spinicauda. —— buccinator. Querquedula crecca. coscoroba. Metopiana peposaca. Dendrocygna autumnalis. Fuligula cristata. viduata. Mergus castor. —— fulwa. Mergellus albellus. The Columbe possess the femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciee; the accessory femoro-caudal is present in all but Lopholemus antareticus. The ambiens may be present or absent; and this feature is probably a subfamily character. ON CERTAIN MUSCLES OF BIRDS. 201 Fig. 5. pp Bi, TN Outer view x of right thigh of Bernicla brenta, partly dissected. s, sartorius; oe, obturator externus; 4d, adductor; 5, biceps; fc, femoro-caudal, afc, accessory femoro-caudal; st, semitendinosus; sm, semimembranosus ; P, pubis ; fv, femoral vein. F The ambiens is present in :— Pterocles alchata. Chamepelia talpacott. arenaria. Metriopelia melanoptera. Lopholemus antarcticus. Tympanistria bicolor. Columba enas. Leptoptila jamaiéensis. livia. Chalcophaps chrysochlora. —— leucocephala. Ocyphaps lophotes. —— picazuro. Phaps chalcoptera. —— maculosa. -Carpophaga globicera. —— vinacea. —— enea. Turtur senegalensis. Calenas nicobarica. — aldabranus. Diduneulus strigirostris. Chaleopelia chalcospilos The ambiens is absent in :— : Goura coronata. -Geopelia striata. —— victoria. —— placida. Page 640. 202 ON CERTAIN MUSCLES OF BIRDS. Geopelia cuneata. Treron calva. humeralis. Phloganas cruentata, Ptilopus melanocephalus. - Starnenas cyanocephala. marie. The Galline possess the ambiens, the accessory femoro-caudal, the semitendinosus (large), the accessory semitendinosus (large), and the postacetabular portion of the tensor fasciz (large); the femoro-caudal is well developed in some, small in some, and absent in afew. An asterisk is placed by the name of those in which it is absent. Species examined. TETRAONIDZ. Tetrao urogallus. tetran. PHASIANIDA. Francolinus vulgaris. afer. ponticerianus. —- gularis. clappertonit. Arboricola torqueola. Perdiz cinerea. Coturniz communis. Rollulus coronatus. Odontophorus dentatus. Ortyx virginianus. Eupsychortyz cristatus. Caccabis chukar. Phasianus colchicus. versicolor. reevestt. Thaumalea picta. amherstice. Luplocamus erythrophthalmus. — vieilloti. —— pyronotus. —— horsfieldit. —— albo-cristatus. ‘ Gallus bankiva. Ceriornis temminckir. * Pavo nigripennis. * muticus. Argus giganteus. * Meleagris gallopavo. Numida meleagris. CRACID2. Craw globicera. —— incommoda. Pauwis mitu. Penelope cristata. pileata. Ortalis albiventris. MEGAPODIDA. Talegalla lathami. Megacephalon maleo. Of the Alectorides the families will be considered separately. In the Bustards as represented by Otis macqueeni and Hupodotis denhami the ambiens, accessory femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz are present; the femoro-caudal is absent. In the Gdicnemide the accessory femoro-caudal, the semitendi- nosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz are present. In Wdicnemus bistriatus and C2. super- ON CERTAIN MUSCLES OF BIRDS. 203 ciliaris the femoro-caudal is present, but very slender. In @. grallarius it is absent. The ambiens is present in the three above-named species, but it is peculiar in being sometimes but imperfectly developed. It _erosses the knee very externally, and was quite normally developed in a specimen of @. bistriatus and in one of @. grallarius. In another specimen of . grallarius it was lost anteriorly on the fasciz covering the knee; and in one of @. superciliaris it crossed the knee, but sent off a slip on the in- and outside to join the investing fascia. In the Cariamide the semitendinosus, the accessory semiten- dinosus, and the postacetabular portion of the tensor fasciz are present. The ambiens was absent in a specimen of Cariama cris- tata, but present in one of Chunga burmeisterit. The femoro-caudai was absent in both. The accessory femoro-caudal was present in Cariama cristata, though extremely small. It was absent in Chunga Among the Gruide, Grus antigone possesses the ambiens, the femoro-caudal (very small), the accessory femoro-caudal (small), the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz, as does also Anthropoides virgo. Among the Eurypygide, Rhinochetus jubatus possesses the ambiens, the femoro-caudal (extremely thin), the semitendinosus, and the _ accessory semitendinosus ; the accessory femoro-caudal is absent. The Rallide possess the ambiens, the femoro-caudal, the accessory femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz. Species examined. Rallus aquaticus. - Ocydromus sylvestris. Page 641. Aramides cayennensis. Porphyrio madagascariensis. Porzana americana. melanotus. carolinensis. Gallinula chloropus. Crez pratensis. The Gralle possess the ambiens, the femoro-caudal, the semiten- dinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascis ; the accessory femoro-caudal may be present or absent. It is present in :— Glareola, sp. Charadrius hiaticula. Numentus arquatus. r Himantopus nigricollis. —— pheopus. Parra africana. Hematopus niger. It is sometimes present (very small), sometimes absent, in :— Charadrius pluvialis. Vanellus cristatus. Page 642. 204 ON CERTAIN MUSCLES OF BIRDS. It is absent in :-— Strepsilas interpres. Scolopax rusticula. Inmosa rufa. Gailinago scolopacina. Totanus calidris. gallinula: solitarius. ~ Calidris canutus. Gambetta flavipes. Tringa cinelus. Machetes pugnaz. The Gavie possess the ambiens, the femoro-caudal, the semiten- dinosus, and the accessory semitendinosus; the postacetabular portion of the tensor fascie is slightly developed in Larus and Lestris, but not in Sterna; the accessory femoro-caudal is present, though small, in Sterna, but absent in Larus and Lestris. Species examined. Lestris antarcticus. Larus glaucus. Larus argentatus. Sterna hirundo. The Procellariide must be considered under two Civanes, the Storm-Petrels and the true Petrels. The Storm-Petrels possess the femoro-caudal, the accessory femoro-caudal, the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fascie#; the ambiens were present in a black one, but quite absent in three specimens of a light-grey species. Species examined. Procellaria pelagica ?* Procellaria fregata ?+ The true Petrels possess the ambiens, the femoro-caudal, and the semitendinosus ; the postacetabular portion of the tensor fascise is very small, or absent, and the accessory femoro-caudal is present in all but Bulweria bulwert. [The accessory semitendinosus is absent. ]} Species examined. GZstrelata lessoni ? Bulweria bulweri. Prion vittata. Puffinus anglorum. Daption capensis. and some unnamed specimens. * [This is, in all probability, Oceanites oceanica. The true Procellaria pelagica lacks the accessory semitendinosus muscle.—ED. ] + [An uncertain species. —ED.] t [Lhis passage has been corrected ; “semitendinosus” (on p. 642) being a mis- print for “ femoro-caudal,” and the absence of the accessory semitendinosus muscle in the birds omitted in the original.—Eb. ] ON CERTAIN MUSCLES OF BIRDS. 205 The Pygopodes consist of two distinct families. Among the Oolymbide, Colymbus glacialis possesses the ambiens, the femoro-caudal, the accessory femoro-caudal, the semitendinosus, and the postacetabular portion of the tensor fascise; the accessory semitendinosus is absent. The Podicipide possess the accessory femoro-caudal, the semiten- dinosus, and the postacetabular portion of the tensor fascie; the ambiens, the femoro-caudal, and the accessory semitendinosus are absent. In these birds the semimembranosus is very thin in Podiceps minor; in specimens preserved in spirit I have not seen it, probably because it is so slender. Species examined. Podiceps cristatus. Podiceps nove-hollandie. — minor. The Alcide possess the femoro-caudal, the accessory femoro- caudal, the semitendinosus, and the postacetabular portion of the tensor fascie; the ambiens and the accessory semitendinosus are absent. Species examined. Alca torda. Arctica alle. Uria troile. The Impennes possess the ambiens, the femoro-caudal, the acces- sory femoro-caudal, and the semitendinosus; the accessory semiten- dinosus and the postacetabular portion of the tensor fasciz are absent. In Eudyptes the insertion of the femoro-candal is much nearer the condyloid end of the femur than to the head. Species examined. Aptenodytes pennantit. Spheniscus humboldti. Spheniscus demersus. Eudyptes catarractes. The Crypturi possess the ambiens, the femoro-caudal, the acces- sory femoro-caudal (which has a slip arising above the sciatic foramen, found elsewhere only in the Struthiones), the semitendinosus, the accessory semitendinosus, and the postacetabular portion of the tensor fasciz. Species examined. Rhynchotus rufescens. Crypturus sallei. Crypturus obsoletus. noctivagus. The Struthiones present marked peculiarities. Page 643. In Apteryz the postacetabular portion of the tensor fascie is very — large, and slightly overlapped at the lowest point of its origin by the 206 — ON CERTAIN MUSCLES OF BIRDS. posterior superior portion of the semitendinosus. The semitendinosus is well developed, and has a broad accessory head. The accessory femoro-caudal is peculiar; for, on removing the biceps cruris, its superficial portion is seen running obliquely upwards and forwards to the whole length of the linea aspera of the femur, from its usual Fig. 6. Outer view of thigh of Apteryx owenii, dissected. s, sartorius ; ¢f, tensor fascie ; b o and b i, biceps, origin and insertion ; gi, I and Iv, gluteus primus and-quartus; s¢, semitendinosus; @ s ¢, accessory semiten- dinosus ; sm, semimembranosus ; sfc, superficial femoro-caudal ; f c, femoro- caudal; a f ¢, accessory femoro-caudal; Ad, adductor. The asterisk on the semimembranosus is on the spot at which its second.or true insertion is; the other asterisk is placed on the slip of the accessory femoro-caudal (in this case specially modified), which is found in Struthious birds and the Tinamous only, _ above the sciatic vessel and nerve. origin. The sciatic artery and nerve are superficial to this muscle (adductor longus of Owen), and parallel to its insertion, as in most birds; but they, as is not the case except in the Struthiones and Crypturi, perforate it at the sciatic notch, leaving a small portion of the muscle (the adductor brevis of Owen) above them. The anterior Page 644. terminal fibres of this muscle are situated external or superficial to the accessory semitendinosus. After this muscle has been removed or turned back, there is seen a ON CERTAIN MUSCLES OF BIRDS. 207 deeper muscle, which, if the one described above did not exist, would be justly considered to be the femoro-caudal and the accessory femoro- caudal, part springing from the iliac ridge and part from the coccyx, whilst both are inserted into the posterior portion of the linea aspera and have the nerve to the semimembranosus situated between them and the adductor magnus. The semimembranosus is also peculiar in having a second head of origin from the ischium, behind the femoro-caudal, and just in front of the origin of the semitendinosus, so that the femoro-candal runs partly in a separate canal between the two heads of origin of this muscle and covered by it. The other muscles are present as in most birds. The ambiens is strong. Species examined. Apteryz owenit. Apteryz mantelli. In Casuarius the postacetabular portion of the tensor fascie is very large, and is overlapped below and behind by the semitendinosus near its origin; the femoral-caudal forms a small cylindroidal belly, which is continued upwards as a narrow tendon; the accessory femoro- caudal is enormous, being perforated by the sciatic artery and nerve ; - it replaces to a great extent the obturator externus, which is pecu- liarly small, and situated anterior to it in origin, as usual. The semi- tendinosus is present and has a broad accessory head. The ambiens is absent. The other muscles are as in most birds. Species examined. Casuarius bennettii. Casuarius bicarunculatus. — galeatus. In Dromeus nove-hollandie the semitendinosus is larger than in Casuarius, and the accessory head is large; the femoro-candal is absent; and the accessory femoro-caudal is very large, being pierced by the sciatic artery and nerve. The biceps cruris is very peculiar in ‘not being inserted in its usual characteristic manner, but ending a little anterior to the middle of the thigh very indefinitely, blending with the fascie in that region, and not being continued directly to the fibula at all. The semimembranosus also is peculiar in having an aponeurotic connection with the middle of the linea aspera, from about the middle of its course. The ambiens is absent. In Rhea americana a somewhat similar condition is found. The biceps is normal; and the semitendinosus, the accessory semiten- dinosus, as well as the postacetabular portion of the tensor fasciz are © much as in Casuarius and Dromeus. The femoro-caudal is absent; Page 111. 208 ON CERTAIN MUSCLES OF BIRDS. the accessory femoro-caudal is large, being perforated by the sciatic artery and nerve ; and the ambiens is strong. In Struthio camelus the ambiens is also well developed. Part II.* (Plate V.) The facts contained in the former part of this communication (“‘ Proceedings of the Zoological Society,” 1873, p. 626) being in an expanded form, it is not easy to appreciate their full significance at a glance, nor without considerable difficulty. To obviate this incon- venience I have constructed the following table, which is so arranged that by a-very simple method, it is possible to tell without further reference which of the five muscles—the ambiens, the femoro-caudal, the accessory femoro-caudal, the semitendinosus, and the accessory semitendinosus—are present or absent. To obtain this result the names of the muscles themselves have been omitted, and single letters of the alphabet used in their stead. The femoro-caudal is represented by.............. A The accessory femoro-candal.................... B The semitendinasns 4:9... f. + on c- Old-W orld. Division @. The oil-gland and gall-bladder present. Genus Oarpophaga. ‘Division y. The accessory femoro-caudal muscle absent (it being present in all other Columbe); the oil-gland and gall-bladder present. Genus Lopholemus: Division 6. The oil-gland and gall-bladder absent. Genus Didunculus. Subfamily Treronivz. Columbide wanting the ambiens muscle. Division « With intestinal ceeca and an oil-gland; no gall- bladder. Genus Phlogenas. NOTES ON TWO PIGEONS. - 241 Division f. With intestinal ceca, no gall-bladder, and no oil- gland. Genus Starnenas. Division y. With an oil-gland, no gall-bladder, and no intes- tinal ceeca. Genus Geopelia. Division 6. With no intestinal cxca, no oil-gland (or a very small one), and scutellated tarsi. Genus Treron. » Ptilopus. Division «. With no intestinal ceca, no oil-gland, no gall- bladder, and tarsi reticulate. Genus Goura. Family Preroctipz. Columbe in which the intestinal ceca con- siderably exceed half an inch in length. Subfamily Prerocuixz. Pteroclide possessing the ambiens muscle, a gall-bladder, and an oil-gland. Genus Pterocles. » Syrrhaptes. 36. NOTES ON TWO PIGEONS, JANTHGNAS LEUCO- LAIMA AND ERYTHRG@NAS PULCHERRIMA-* SINcE my communication to the Society “ On some Points in the Page 367. Anatomy of the Columbe,”* specimens of two species of this group have died in the Gardens, which deserve a passing note. Tantheenas leucolema.—The genus to which this bird belongs has been, by different authors, placed sometimes in the Columbine and at others in the Carpophagine-section of the family—the number of the rectrices (12, and not 14) having made its position uncertain, as its general appearance tends to that of the Fruit-eaters. From the paper above referred to, the definition of the Columbine, Page 368. containing the genus Columba, may be thus stated :— Cotumsinz. Columbide possessing an ambiens muscle, intestinal ~ cxeca, an oil-gland, 12 rectrices, and no gall-bladder. . *® “Proceedings of the Zoological Society,’ 1875, pp. 367,8. Read, May 4, 1875. ; + “ Proceedings of the Zoological Society,” 1874, p. 249 et seqq. Page 471, 242 ON THE AUSTRALIAN BUSTARD. Whereas Carpophaga possesses the ambiens muscle, an oil-gland, a gall-bladder, and no intestinal ceca. In Ianthenas leucolema the ambiens muscle and the oil-gland are present, as are the intestinal ceeca.* The gall-bladder isabsent. This bird must therefore, together with Columba, Turtur, Macropygia, and Ectopistes, be placed in the Columbine and not in the Carpopha- gine division. The intestines are 47 inches in length, of average diameter; and the gizzard is typical in structure, having simple plicated pads. Erythrenas pulcherrima.—This species is truly Ptilopine in all its characters. As in Ptilopus, the ambiens muscle is wanting, as are the ceca to the intestine. The gall-bladder is present; and the oil-gland is very small. The gizzard presents the peculiarities of that genus, although the four pads are not so regularly constructed, minor plications existing. There are 14 rectrices; and the intestines (which are capacious, as in all fruit-eating birds) are 16 inches in length. 37. ON THE “SHOWING-OFF” OF THE AUSTRALIAN BUSTARD (EUPODOTIS AUSTRALIS). WuetHER the account of the production of great distention of the neck in the male Australian Bustard which follows will in any way simplify the question of the presence or absence of a gular pouch in Bustards generally, is doubtful. At all events it will rectify an ac- cepted error, and add a fresh fact to the considerable literature of the subject. In the “ Proceedings” of this Society for 1868 (p. 471 et seq.), Dr. Murie pictures the sexual “ show off” in a specimen of Hupodotis australis which was presented to the Society in April 1866, by the Acclimatization Society of Sydney, and infers, from its appearance, that, as an undoubted fact, the gular pouch is present in this specimen of the species at least. In 1873, during one of the months in which it was “ showing off,” namely in May, 1 examined the mouth of this identical bird while * These are extremely slender, and require special precautions to be taken for their demonstration. + “Proceedings of the Zoological Society,” 1874, pp. 471-3. Read, June 16, 1874. ON THE AUSTRALIAN BUSTARD. 243 alive, and could find no trace of a sublingual orifice, and, what is more, felt and saw a median frenum lingue quite distinctly. This made me doubt the correctness of Dr. Murie’s inference, that, because Page 472. the neck of Eupodotis australis becomes distended much during the sexual season, therefore there is a gular pouch. This individual bird, which formed the subject of Dr. Murie’s plate (“ Proceedings of the Zoological Society,” 1868, pl. xxxvi.), died on May 11, 1874, having shown off in its wonted manner during the few preceding weeks. An excellent opportunity was thus afforded for the decision of the question whether or not this specimen had a gular pouch. Fig. 1. Fig. 2. Fig. 1. The esophagus, trachea, and gular pouch of a specimen of Otis tarda, seen from the side. The crop is here ps8 rawn as in the actual preparation, projecting backwards, and not forwards as usual. Fig. 2. The esophagus and trachea of the specimen of Eupodotis australis here described. The wsophagus is much dilated, and, like that of the Pouter Pigeon, can be distended with air by the living bird. No trace of a pouch or crop is to be seen. There was no gular pouch. There was no sublingual orifice. The freenum lingue was well developed, it being necessarily quite absent in the adult male of Otis tarda. How unsafe therefore is it to infer that, because the neck distends and depends during the “ show-off,” there R 2 Page 473. 244 ON THE AUSTRALIAN BUSTARD. must be a sublingual pouch. It is quite possible that two effects, very similar in appearance, in closely allied birds, may be the result of different mechanisms. In the feet of the Cuculide and the Picide the scansorial arrangement of the toes is the result of entirely different dispositions of the tendons which move them; and in Otis tarda and Eupodotis australis the same reasoning holds. In both these birds there is, during the show-off, a distention with air of a well differentiated bag, which is in both cases lined with a true mucous membrane. But in Otis tarda this sac is a special struc- ture in front of the windpipe, opening under the tongue; whilst in Eupodotis australis (in the specimen under consideration at least), it is simply a highly dilated cesophagus. _ Through the kindness of Lord Lilford I am in the possession of an excellent Spanish specimen of the gular pouch of Otis tarda (see fig. 1, p. 243), with the whole of the cesophagus, the tongue, and part- of the trachea attached. In it the gular pouch, opening sublingually, is capacious, and, when distended, egg-shaped with no constriction in any part. The csophagus is uniformly cylindrical for its upper two thirds, and not at all enlarged. Lower down there is a well- developed globular crop. In the specimen of Hupodotis australis which died on May 11, as previously mentioned, there is no trace of a gular pouch. The eso- phagus is enormously dilated from its commencement (see fig. 2, p- 243), and gives no indication whatever of any division into tube and crop. Its greatest cireumference, when fairly inflated, is 14 inches, and the length of the distended portion of the tube is 174 inches. Before dissection, by filling its cavity with air, the lower portion of the dilated cesophagus protruded downwards considerably in front of the symphysis furcule, and formed the depending portion of the sac which was so conspicuous in the living animal. The trachea de- scended in front of this sac ; and when the latter was undistended, the former, on account of the diminished distance between the points it had to reach, was zigzagged from side to side in the part opposite the pendant portion. The keeper, J. Church, tells me that, when handling the sac in the living bird, he always felt a hard cord running down in front of it, which was evidently the windpipe. The dilated cesophagus was, as might have been expected, covered with two coats of muscular tissue, the outer longitudinal—and the inner transverse. The mucous lining presented no peculiarities. The skin in front of the neck was lax, with a considerable amount of coarse fat in its deeper layer ; it was engorged with blood, tortuous vessels running through it in all directions. I may mention as an anatomical peculiarity of interest that Huwpo- dotis australis and EH. denhami possess but one carotid artery, the right ON THE “ SHOW-OF” IN BUSTARDS. 245 —a condition I have not seen in any other bird; Ofis tarda and O. macqueent have two, and Tetraz campestris the left only. Most pro- bably the presence of a right carotid only is charaoteristic of the genus Eupodotis. a FURTHER NOTE ON THE MECHANISM OF THE « SHOW-OFF ” IN BUSTARDS.* Ir is the uncertainty with which my material comes to hand which Page 673. must be my excuse for having so soon to present a further note on the “ show-off ” in the Bustards. A young male specimen of the Great Bustard (Otis tarda) has recently died in the Society’s Gardens; and one or two observations which I was able to make on its gular arrangements have done much to clear up, in my mind, the difficulties connected with that some- what involved subject. My previous communication on this point (‘‘ Proceedings of the Zoological Society,” 1874, p. 471)+ contains a ~ drawing of the esophagus, trachea, and gular pouch of a Spanish specimen of Ofis tarda, kindly given me by Lord Lilford. In the description appended to the woodcut it is remarked that the crop is peculiar, in that it springs from the posterior instead of the anterior wall of the cesophagus ; and I may mention that it is further peculiar in not being quite median, as would have been expected. I do not know the age of the young male bird above referred to, - which I have recently examined. It seemed of nearly full size, had been in the possession of the Society between three and four months, Page 674. had never shown off, and had no lateral tuft of feathers from the sides of the lower jaw. In it the esophagus was uniformly cylindrical, with no trace of a crop, and there was no gular pouch. On looking under the tongue, however, it was evident that the arrangement of the sublingual struc- tures was quite peculiar. In the male of Eupodotis australis, as I have previously remarked,} the frenum lingue is well developed in the normal manner as a median vertical fold; and, what is more, it is situated as far forward as in most animals, not behind the level of the * “Proceedings of the Zoological Society,” 1874, pp. 673, 4. Read, Dec. 1, 1874. + (Supra, p. 242.) } “ Proceedings of the Zoological Society,” 1874, p. 472. (Opp.) piel getre i: 246 ON THE “ SHOW-OF” IN BUSTARDS. basihyal apparatus. In the young and pouchless male of Otis tarda the condition is very different. In it the frenum lingue does not exist as such, but as two slight lateral vertical folds, with a median interval between them, a quarter of an inch across; so that the pouchless sublingual region of the young male Otis tarda is very like the excel- lent drawing of that of the pouched adult male in Dr. Murie’s paper on the bird (“ Proceedings of the Zoological Society,’ 1869, p. 141), except that what is there represented as an aperture to a pouch must be considered for the time being as only a slight depression. The tongue is also free for a considerably further distance along its under surface than in Eupodotis australis. In a specimen of the head of Otis tarda in the Museum of the College of Surgeons* the freenum lingus is median and normal in all respects. The sex is not mentioned; but from the fact of its differ- ing so much from that of my young male specimen, I cannot help inferring that it is that of a female. If such is the case, until more examples are obtainable, the certainty as to the correctness of my surmise is not absolute. — The two sublingual frena, with a membrane between them, make it seem almost certain to me that in the adolescent male bird, and not in the female, there is every opportunity for the development of a pouch, and that the habit of inflating the air-passages during the sexual season distends the membrane between the frena lingue, it being comparatively weak, and causes it to develop into a pouch from continued stretching. In favour of the here assumed existence of considerable pressure is the existence of the abnormally situated di- verticulum in the specimen figured in my previous paper on the subject; for, from the absence of any trace of a crop in the young bird, it may be inferred that such an organ does not pertain to the species; therefore it must be the result of some superadded force, brought into action in the adult, the distention of the pharynx during the “show-off” being quite sufficient to account for it. The specimen figured in my earlier communication and that described in the present may all be seen in the Museum of the College of Surgeons. * No. 772 Q. ON THE ANATOMY OF THE PARROTS. 247 39. ON SOME POINTS IN THE ANATOMY OF THE PARROTS WHICH BEAR ON THE CLASSIFICA- TION OF THE SUBORDER.* (Plates VI & VIL.) Iy a former communicationt, a review of certain of the most variable Page 586. characters found amongst the Columbe enabled me to give hints with regard to the mutual relationships of the different genera of that con- siderable family, which I hope will be found of service. On the present occasion it is my desire to follow out a similar method, taking the Psittaci, a suborder quite as, and perhaps even more, difficult to by external features only. The unequalled collection of living Parrots in the Society’s Gardens, Page 587. and the liberality of friends, have placed at my disposal specimens ' of the large number of 82 species, in all of which I have been able to note the characters laid stress on in this paper. To save repetition a list is here given of the species examined by me; and on all future occasions when a genus only is mentioned, it refers to those species of it which are contained in this list. Names of Species examined. Agapornis roseicapilla. Caica melanocephala. Aprosmictus erythropterus. - Calopsitia nove-hollandie. scapulatus. Calyptorhynchus banksit. Ara ambigua. Chrysotis agilis. ararauna. collaria. —— macao. : —— festiva. maracanda. — levaillantit. Bolborhynchus monachus. ochrocephala. Brotogerys tiriacula. ; Conurus eruginosus. —— tovi. aureus. -—— pyrrhopterus. —— aztec. virescens. —— hematorrhous. Cacatua cristata. —— holochlorus. —— galerita. —— jendaya. —— leadbeateri. —— monachus. —— sulphurea. —— nanday. * “ Proceedings of the Zoological Society,” 1874, pp. 586-98. Pls. LXX and t “Proceedings of the Zoological Society,” 1874, p. 249. (Supra, p. 229.) Page 588. 248 Conurus pavua. peta. xvantholemus. Ooracopsis barklyi. ' Cyanorhamphus auriceps. nove-zealandic. Eclectus polychlorus. Holophus roseicapillus. Hos cardinalis. —- indica. riciniata. Euphema aurantia. bourkit. — pulchella. splendida. Geopsittacus occidentalis. Lathamus discolor. Licmetis pastinator. Loriculus asiaticus. chrysonotus. galgulus. Lorius lort. tricolor. Melopsittacus undulatus. Nestor meridionalis. ON THE ANATOMY OF THE PARROTS. Nestor notabilis. Paleornis alexandri. erythrogenys. longicauda. torquata. Pionus menstruus. sordidus. Platycercus eximius. pallidiceps. zonartus. Peocephalus fuscicapillus. senegalensis. Prioniturus, sp. Psephotus hematogaster. pulcherrimus. Psittacus erithacus. Psittacula passerina. Psittinus malaccensis. Pyrrhulopsis splendens. Pyrrhura leucotis. vittata. Stringops habroptilus. — Tanygnathus muelleri. Trichoglossus concinnus. novee-hollandice. The points to which my attention has been directed, on account of the variations observed, are :— 1. The arrangement of the carotid arteries ; 2. The presence or absence of the ambiens muscle ; 3. The presence or absence of the furcula; 4, The presence or absence of the oil-gland ; and others of minor importance, such as the complete encirclement of the orbit by bone, and the peculiarities of the atlas vertebra. These will be considered separately. I. The arrangement of the carotid arteries among the Parrots. In my paper on the carotid arteries of birds, the peculiarities of these vessels in the Parrots are described, it being shown that three different arrangements of these vessels obtain, and perhaps a fourth. Hither the two carotids may run normally, independent, and side by side up the front of the neck, in the hypapophysial canal ; or the right, as usual, traverses that canal, whilst the left rans superficially along ON THE ANATOMY OF THE PARROTS. 249 the side of the neck in company with the left pneumogastric nerve and the jugular vein; or the left carotid may alone be developed, as in the Passeres and many other birds. It has been stated that Meckel found _@ fourth arrangement in Cacatua sulphurea; but in a specimen of that species recently dissected by me, the left only was present, as in C. cristata, C. leadbeateri, and C. galerita. To make this paper complete in itself, and to incorporate those dissections performed since the other was published, a list of all the Parrots which I have examined, together with the condition observed in them is given. 1. I eecclidirdae peceentt, rasinitgy niosmallf, eidd BY side, im Agapornis, Loriculus, Aprosmictus, Lorius, Calopsitta, Melopsitiacus, Calyptorhynchus, Paleornis, Eeclectus, Prioniturus, Eolophus, Psittinus, Eos, Stringops, Euphema, Tanygnathus, Geopsittacus, Trichoglossus. Liemetis, é 2. The left carotid only is present in Cacatua. 3. The two carotids are present, the right having its normal _ course, the left running superficially along with the left pneumogastric nerve in Ara, Nestor, Bolborhynchus, Pionus, Brotogerys, Platycercus, Caica, Peocephalus, Chrysotis, | —- Psephotus, Conurus, Psitiacus, Coracopsts, Psittacula, Cyanorhamphus, _ Pyrrhulopsis, Lathamus, Pyrrhura. Page 589. It may be observed that the only other well-defined groups of birds in which the carotids vary are the Cypselide, Gallinz, Struthiones, and Otidex. ~ Il. The presence or absence of the ambiens muscle. The ambiens muscle, the tendon of which crosses the front of the knee-capsule obliqnely from above downwards and outwards, and ulti- Taal . fe Page 590. 250 ON THE ANATOMY OF THE PARROTS. mately forms part of the flexor perforans digitorum, is present in the following genera— Ara, Nestor, Bolborhynchus, Peocephalus, Caica, Psittacus, Conurus, Stringops. It is absent in Agapornis, Licmetis, Aprosmictus, Loriculus, Brotogerys, Lortus, Cacatua, Melopsittacus, Calopsitta, Paleornis, Calyptorhynchus, Pionus, Chrysotis, Platycercus, Coracopsis, Prioniturus, Cyanorhamphus, Psephotus, Eclectus, Psittacula, Holophus, Psittinus, Eos, Pyrrhulopsis, Euphema, Pyrrhura, Geopsittacus, Tanygnathus, Lathamus, Trichoglossus. The only other well-defined groups of birds in which the ambiens muscle is known to vary are the Columbe and the Struthiones. TII. The presence or absence of the furcula. By this expression is meant the presence or absence of the furcula as a complete bone; for in those Parrots in which it is said to be absent, the scapular ends of the two parts of which it is composed are frequently to be found, being of considerable length in Stringops and some of its allies. The furcula is complete in Aprosmictus, Coracopsis, Ara, Eclectus, Bolborhynchus, Holophus, Brotogerys, Hos, Cacatua, Lathamus, Caica, Tiemetis, Calopsitta, Loriculus, Calyptorhynchus, Lorius, Chrysotis, Nestor, Conurus, Paleornis, _—_— ON THE ANATOMY OF THE PARROTS. Pionus, Peeocephalus, Prioniturus, Psittacus, Psittinus, Agapornis, Cyanorhamphus, Euphema, Geopsittacus, Melopsittacus, Pyrrhulopsis, Pyrrhura, Tanygnathus, Trichoglossus. The furcula is but partially developed or absent in Platycercus, Psephotus, Psittacula, Stringops. 251 Dr. Finsch, in his monograph on the Parrots*, has given a list of IV. The presence or absence of the oil-gland. those species in which the condition of the furcula has been recorded, which is very complete, embracing most of the above genera. When present the oil-gland is always tufted in the Parrots. It is present in the following genera :— . Agapornis, Aprosmictus, Ara, Bolborhynchus, Caica, Calopsitta, Calyptorhynchus, Conurus, Coracopsis, Cyanorhamphus, Eclectus, Eolophus, Eos, Euphema, Geopsittacus, Lathamus, LTiemetis, It is absentin - Brotogerys, Loriculus, Lorius, Melopsittacus, Nestor, Paleornis, Platycercus, Peeocephalus, Prioniturus, Psephotus, Psittacula, Psittacus, Psittinus, Pyrrhulopsis, Pyrrhura, Stringops, Tanygnathus, Trichoglossus. Pionus. Nitzsch, in his work on pterylosist, has recorded its absence in some of the genera. * “Die Papageien,” Bd. i. p. 197. t+ “ Pterylography,” English edition, p. 98 e¢ seq. Page 591. 252 ON THE ANATOMY OF THE PARROTS. In Cacatua galerita and C. leadbeateri it is present; but it is generally wanting in OC. cristata, and has not been found in C. sul- phurea. The above facts may be tabulated in a form which makes their significance more readily apparent, by placing those together in which a similar arrangement is observable. Thus there are :— 1. Parrots in which there are two normally situated carotids, no ambiens muscle, a furcula, and an oil-gland—namely : Aprosmictus, Loriculus, Calopsitta, Lorius, Calyptorhynchus, Paleornis, Eclectus, Prioniturus, Holophus, Psittinus, - Eos, Tanygnathus, Liemetis, Trichoglossus. 2. Parrots in which there are two normally situated carotids, no ambiens muscle, no furcula, and an oil-gland—namely Agapornis, Geopsittacus, EHuphema, Melopsittacus. 3. Parrots with a left carotid only, no ambiens muscle, a furcula, and generally an oil-gland—namely ; Cacatua. 4. Parrots with two carotids (the left being superficial), an ambiens muscle, a furcula, and an oil-gland—namely Ara, Nestor, Bolborhynchus, Paocephalus, Caica, Psittacus. Conurus, 5. Parrots with two carotids (the left being superficial), no ambiens muscle, a furcula, and an oil-gland—namely Coracopsis, Pyrrhulopsis, Lathamus, Pyrrhura. 6. Parrots with two carotids (the left being superficial), no ambiens muscle, a furcula, and no oil-gland—namely Brotogerys, Pionus. Chrysotis, 7. Parrots with two carotids (the left being superficial), no ambiens muscle, no furcula, and an oil-gland—namely Cyanorhamphus, Psephotus, Platycercus, Psitiacula. ON THE ANATOMY OF THE PARROTS. 253 The true significance of these facts next requires attention; and the principle upon which all attempts at the formation of a satisfac- tory genealogy or classification of the suborder can be arrived at must be born in mind throughout. It is the following:—An anatomical Page 592. character is so much the more or less certain to have been an element of the original type or ancestor whence sprang the class, order, family, or genus under consideration as it is more or less frequently found in the less intimately related minor divisions of the groups under obser- vation. An example will make this more clear :—T wo large arteries (the carotids), one on each side, run up to supply the head in most Pulmonate Vertebrata, as faras I know. In all Mammalia such is certainly the case. In many Birds there are, similarly, two carotids, though some have only one. It is therefore more than probable that the ancestral bird had two carotids, those in which one is absent having lost it subsequently. Many Parrots have two carotids; the genus Cacatua is characterised by the left only being present: it, therefore, has in this respect departed most from the ancestral type. Again, other Vertebrata and other Birds with both carotid arteries present have them symmetrically placed; many Parrots have symme- trical carotids; but in some the left (and the left only) is abnormal in being superficial: therefore, from the same considerations, these last have differentiated off from the parent stem, and, what is more, this ’ peculiarity can hardly have occurred on more than one occasion, as it is otherwise unique and therefore peculiar and exceptional in origin. There is another principle to be remembered, which is that there is no such thing as reversion to lost ancestral anatomical characters. The genus Cacatua has lost its right carotid, as have the whole family of the Passeres and many others. There is not a tittle of evidence in favour of the assumption that they or their descendants could ever regain that vessel. Its arrested development is a positive act, the result of extra forces coming into play in early embryonic life, to remove which would require the introduction of a certain definite geries of counterbalancing forces superadded to those already in action; whilst in the ancestral bird, the persistence of the two arteries resulted from the absence of any impediment to their deve- lopment. The probability that the ancestral form should be reverted to cannot be greater than that an entirely new arrangement should be effected. That some domestic excentric varieties should tend in some cases to revert to the wild type can have no more bearing on the general subject than the similar tendency to exaggeration which is not apparent in the feral forms. Upon these principles many deductions can be made as to the mutual relations of the several genera of the Psittacine suborder. For instance, it must be inferred that the ancestral Parrot possessed P ¢ ha Page 593. 254 ON THE ANATOMY OF THE PARROTS. two carotids, running symmetrically in front of the neck, and that the ambiens muscle was present, as was the furcula and the tufted oil- gland. The intestinal ceca and gall-bladder must have been absent or lost very early, as must the postacetabular portion of the tensor fascize muscle ;* for they are none of them to be found in any existing species; whilst the beak, tongue, crop, and rectrices must have possessed the characteristic features, which are not found to vary to any important extent. The pterylosis of the suborder forms a con- siderable but much involved field for work, which has only been entered upon by the illustrious Nitzsch. Referring back to the characterising features of the existing species whose internal structure has been noted, it will be seen that none has as yet been found with a conformation exactly similar to that of the above-described ancestral bird ; in other words, no existing Parrot has been seen with two normal carotids, an ambiens muscle, a furcula, and an oil-gland. By more than a single way, however, this condition, with only one exceptional character, is found to exist. For instance, the fourth combination above given, in which the ambiens, furcula, and oil-gland are present at the same time that the carotids are abnormal (the left being superficial), agrees with the type except in one point—the disposition of the carotid arteries. Again, in the first of the combinations the only deviation from the type consists in the absence of the ambiens muscle. These two different directions of variation must therefore have formed the secondary stems from which the more specialized genera subsequently sprang. In other words, the main stem must have given rise to two, in one of which the carotids remained normal, whilst in the other the left became superficial. The following are the genera as they will thus appear :— Genera in which the left carotid Genera in which the left carotid has remained normal. has become superficial. (PALHORNITHIDZ. ) (PsI?TaciDzZ. ) Agapornis. Ara. Aprosmictus. Bolborhynchus. Cacatua. Brotogerys. Calopsitta. Caica. Calyptorhynchus. Chrysotis. Eclectus. Conurus. Eolophus. Ooracopsis. Eos. Oyanorhamphus. EHuphema. Lathamus. * Vide “ Proceedings of the Zoological Society,” 1873, p. 628. (Supra, p. 189.) a OO OO ———— ON THE ANATOMY OF THE PARROTS. 255 Genera in which the left carotid Genera in which the left carotid has remained normal. has become superficial. (PALZORNITHIDZ) (PsITTAcIDz2.) Geopsittacus. Nestor. Loriculus. Pionus. Lorius. Platycercus. Melopsittacus. - Peeocephalus. Paleornis. Psephotus. Prioniturus. Psitiacus. Psittinus. Psitiacula. Stringops. Pyrrhulopsis. Tanygnathus. Pyrrhura. Trichoglossus. Each of these secondary types must have then become a centre for variation in itself. From the 4th combination, in which only the carotids are peculiar, sprang the 5th, 6th, and 7th, with the ambiens deficient, just in the same way that the 1st, 2nd, and 3rd combinations originated from the ancestral form by the same process of reduction. The loss of the furcula and of the oil-gland (though never, as it Page 594. happens, both together) have further aided in the formation of tertiary and further subdivisions, which, upon the dissections above given, would lead to the arrangement of the family which is depicted in Plate LXX [6.]. This may be represented in the tabular form as follows :— Suborder (or Cohort) PSITTACI. Family I. Panzornituipz. (Left carotid normal.) The ambiens muscle absent. Carotids two, except in the genus Cacatua. Subfamily (1) Panzorxiramnz. No further deviation. Paleornis. Eclectus. Aprosmictus. Tanygnathus. Prioniturus. Psittinus. Loriculus. Trichoglossus. Lorius. Eos. Subfamily (2) Cacarurnz. Orbital ring completely ossified, and characteristic in that it develops a process bridging the temporal fossa (vide Plate LXXI [7.]). 256 ON THE ANATOMY OF THE PARROTS. Calopsitta, Calyptorhynchus. Inemetis. ‘ Eolophus. Cacatua. Subfamily (3) Srrivcorinz. The furcula, lost.* Stringops. Huphema. Geopsittacus. Melopsittacus. Agapornis. Family II. Psrrracipz. (Left carotid superficial.) Division 1. The ambiens muscle present. Subfamily (4) Arinaz. No further deviation. Ara. Oonurus. Bolborhynchus. Oaica. Psittacus. Peocephalus. Nestor. Page 595. Division 2. The ambiens muscle wanting. . Subfamily (5) Prrraurinz. No further deviation. Pyrrhura. Lathamus. Coracopsis. Pyrrhulopsis. Subfamily (6) Piarycercinaz. Furcula lost. Platycercus. | Pett Oyanorhamphus. Psittacula. Subfamily (7) Curysorina. Oil-gland lost. Chrysotis. Pionus. Brotogerys. In phyllogenetic language this arrangement would be expressed thus:—The original stem, in which the carotids were normally dis- * See the special remarks on this group in the postscript to this paper. Se _— ON THE ANATOMY OF THE PARROTS. 257 posed, gave off a branch characterised by their abnormal arrange- ment. The continuation of the main stem, as well as the branch, shortly lost the ambiens muscle—the latter (and not the former), however, being represented at the present day in its unmodified form by the Arinw. Each of the two secondary branches persists as the Paleornithine and the Pyrrhurine respectively, they both branching dichotomously in an exactly similar manner, the former giving rise to the Stringopine and the specially modified Cacatua cristata and C. sulphurea, the other to the similarly modified Platycer- cine and Chrysotine. A method of formulation will assist in making this more readily intelligible. If the presence of two carotids, normally disposed, is represented by the figure 2, the abnormal arrangement may be re- presented by 2. Then, if the presence or absence of the varying structures be represented by + or —, the following table will express the characteristics of the different subfamilies :— P Carotids. | Ambiens. | Furcula. | Oil-gland. Palzornithine 2 - + + . 2 _ = - Cacatuine .. { L = ie s Stringoping ....... 2 oa ~ Aring ...... 2 7 ~ - Pyrrhurine ....... 2 oo - ~ Platycercine....... 2 = _ - Chrysotine .. 2 - - - Type form .. 2 ~ + rs The Paleornithine will thus be represented by the formula 2, —,+, +; the Arine by 2, +, +, +; and soon. Plate [7] LXXI. will represent these facts in a more self-evident manner. Through the kindness of Prof. Flower and also from the death of the specimen presented by Mr. Murdoch to the Society, I have twice Page 596. had the opportunity of dissecting Stringops habroptilus. As a Parrot it is not so strikingly peculiar as many seem to think. Its wings are useless, and the carina sterni is correspondingly reduced, it is true ; but as points of classificational importance, I regard these as insignifi- cant. The points of special anatomical interest which it does possess, however, are particularly instructive. The proximal ends of the incomplete furcula are well developed, so much so that it might at first sight seem that their symphysial ends are only lost in correlation with the excessive reduction of the powers of flight; though this is probably not the case, because the allied similarly modified genera Euphema &c. do not keep to the ground. 3 Further, in the Society’s specimen above mentioned, though the s Fo a Tl a Page 597. 258 ON THE ANATOMY OF THE PARROTS. ambiens muscle did not cross the knee, yet its fleshy belly was well differentiated on both sides, its thin tendon being lost over the capsule of the joint. In the College of Surgeons’ specimen, however, this muscle was entirely absent in the only knee which was in a fit state for dissection, the other being much shot. It is only in the genus CGidicnemus that I have elsewhere found a similar partial loss of the ambiens.* The partial development of this muscle in this particular instance shows that the tendency to lose it is not of great antiquity ; and it is to be noted that there is no other Parrot with norma: carotids in which any trace of an ambiens is to be found. These considera- tions suggest, what may perhaps be the case, as is suggested by the peculiarities of their geographical distribution, that Agapornis may be the representative among the normal-carotid Parrots of the Platycer- cine branch from the Arinz, whilst the Stringopine proper (including Geopsittacus, Melopsittacus, and Huphema) are more direct continua- tions of the main stem, as indicated by the dotted portions of the diagrams (Plates [6] LXX. and [7] LXXI.), Stringops itself being the nearest living representative of the common ancestor of the whole suborder. Further, it may be worth while taking a glance at some of the most important changes which my classification would necessitate. Taking Mr. Sclater’s “Revised List of Vertebrated Animals in the Society’s Gardens” as a good representative of current opinion, the Order Psrrract is there divided into two families and seven subfamilies, thus :— Order PSITTACI. Family I. StRiINGOPIDA. Family II. Psirtacipa. Subfamily 1. Cacarurnz. ro 2. ARINZ. +5 3. PLATYCERCINE. + 4, PsITTAcIna”. if 5. LORIINA. ‘5 6. NESTORINA. As far as the major division is concerned, the facts brought forward in the present communication suggest a different arrange- ment, as shown above, which would approximately distribute these subfamilies thus :— Family I. Family IT. Stringopide. Arine. Loriine. Platycercine (in part). * Vide “ Proceedings of the Zoological Society,” 1873, p. 640. (Supra, p. 202.) ON THE ANATOMY OF THE PARROTS. 259 Family I. Family TH. Cacatuine. Psittacine (in part). Platycercine (in part). Nestorine. Psittacine (in part). The generally received families Platycercine and Psittacine are avowedly rather incongruous mixtures. Mr. Gould, with acute per- ception, was able to differentiate Aprosmictus from Platycercus, chiefly by its habits of life; and anatomical considerations show that Dr. Finsch’s attempt to reabsorb it in the older genus is a retrograde step. If Dr. Meyer is right in stating that the sexes in Hclectus are of different colours, its relations to Aprosmictus may be very intimate. — Tt may at first sight seem very heretical to remove Lathamus from the Loriinz, the brush-tongue being considered characteristic of that subfamily. To the unbiased student, however, the brush-tongue is a character not more important than several of those that have been above considered. It is only an excessive development of the papille which are always present on the lingual surface, and is seen in a slightly different form in the Lion and other Felidw. The character of the papille is somewhat different in Lathamus from what it is in Lorius, they being blunter and shorter in the former genus than in the latter. The totally different geographical distribution of Paleornis and the true Arinz is quite opposed to Dr. Finsch’s proposition that Conurus and Brotogerys should be the neighbours of the Paleogzean genus. Brotogerys entirely agrees in structure with Chrysotis and Pionus, differing greatly from Conurus; whilst in itself Conurus, as gene- rally received, embodies the red-tailed species, with the fourth primary not acuminate, and the green-tailed species, with an accumi- nate fourth primary. In the former section (Pyrrhura) the ambiens muscle is quite lost, whilst in the latter (Conurus) it is always well developed. : Prof. Huxley is not the only naturalist who has been puzzled by the geographical distribution of Psittacula. This genus in its wide sense, however, is broken up into far separated genera—the Old- World Psittinus and Agapornis differing entirely from the New-World Psittacula, Agapornis being the homologue, as it may be termed, in the normal-carotid Parrots of Psittacula in the other group, whilst Psittinus is a less differentiated genus of the former division. ~ Nestor no doubt stands rather isolated ; but possessing the ambiens muscle, as well as all the other characters of Psittacus and the true Arinz, it must be classed with them. My subfamily Pyrrhurine seems a mixture; and all I can say in its favour is that the combination of anatomical characters is exactly Page 598. the same in all its genera, which have a very scattered distribution. s 2 260 ON THE ANATOMY OF THE PARROTS. Tt will be noticed that no Parrots with normal carotids occur in the New World: and, as far as I know, none but members of that group have red beaks. P.S. (December 8th, 1874).—On the 25th of last month, from the death of one of the specimens of Stringops habroptilus, recently pur- chased by the Society, I have had an opportunity of dissecting a. third individual of the species. In it the ambiens muscle is complete, of fair size, at the same time that it crosses the knee as in Psittacus. This makes me feel more convinced that the arrangement indicated by the dotted portions of the diagram plates accompanying this com- munication is the correct one, and that the main stem has given rise to three instead of two branches—the Stringopine being the nearest representatives of the ancestral form, some of its members (Geopsitta- cus, Melopsittacus, Buphema, and Oyanorhamphus) having quite re- cently lost, whilst Stringops itself is just now on the point of losing the ambiens muscle. It is, however, quite possible, if external resemblances and geographical distribution are left out of considera- tion, that Stringops must stand as the sole representative of the Stringopinez, thus conforming with generally received ideas; and that Geopsittacus, together with Melopsittacus, Euphema, and Oyanorham- phus, rust be placed with Agapornis as part of the family Agaporni- thine, in which the formula is 2, —, —, +. The close external resemblance between Stringops and Geopsittacus nevertheless makes me indisposed to adopt this view. ON THE ANATOMY OF CERTAIN PARROTS. 261 40. NOTES ON THE ANATOMY OF CERTAIN PARROTS.* Sryce the publication of my paper “‘On some points in the Anatomy Page 691. of the Parrots,” in the “ Proceedings” of the Society (1874, p. 586), I have had the opportunity of dissecting several species, some of which, from their peculiarities, deserve special note. They are the following :— 1. Dasyptilus pecqueti. It is through the great kindness of Dr. A. B. Meyer that I have had the advantage of being able to dissect this extremely rare bird. Dr. Meyer obtained the specimen in New Guinea, and has preserved it in spirit, entire. He has most liberally allowed me to determine those anatomical points to which I have drawn attention in the paper above referred to. There are two carotid arteries; the left, however, runs superficially as in the Psittacide (as defined by me). The oil-gland is large, sub- globose and possesses a well-developed tuft of shortish feathers around its orifices. The rectrices are twelve in number. In its myology and osteology it agrees with the Pyrrhurine in entirely wanting the ambiens muscle, and in having a furcula, which bone is not large or powerful, nor so slender as in the Lories. The orbital ring is incomplete, the eye not being encircled by bone. The femoro-caudal muscle is large; and the semitendinosus with its ,accessory head are well developed,—in which arrangements it agrees with all the Psittazi. The intestines are 17} inches long, there being no trace of cxca. The liver-lobes are somewhat unequal in size, the left being the smaller. The stomach is small and much like that in the Fruit-eating Parrots generally. The proventriculus forms a dilated sac, of which the walls, instead of being, as is usually the case, thick and glandular, are strikingly thin, at the same time that no glands are visible. Dr. Meyer has alreadyt given a short description of the peculiari- ties of the tongue, and, in writing to me, tells me that he has further observations to make on the same. ; _ As in nearly all birds, the main artery of the thigh is the sciatic, Page 692. : whilst the vein is the femoral. There is a fenestra near the posterior margin of the sternum, on each side. * “Proceedings of the Zoological Society,” 1876, pp. 691-2. Read, June 20, 1876. + “ Mittheil. aus dem k. zoologischen Museum zu Dresden,” 1875, p. 14. 262 ON THE ANATOMY OF CERTAIN PARROTS. 2. Deroptyus accipitrinus. A Brazilian specimen of this rather peculiar genus from the Society’s collection has the two carotids arranged as in Dasyptilus, the left being superficial. The furcula is of fair size, the orbital ring incomplete, the oil-gland well tufted, the ambiens muscle absent. 3. Polyteles barrabandi possesses two carotids, normally situated— in other words, side by side in the hypapophysial canal. The furcula issmalland slender. The oil-gland is decidedly large, and well tufted. There is no ambiens muscle. The intestines measured 40 inches. 4. Chalcopsitta scintillata has the two carotids normally situated, a small furcula,a small tuft to the large oil-gland, and no ambiens muscle. The intestines measured 37 inches. 5. Coriphilus fringillaceus has the two carotids normal, the furcula small and slender, no ambiens muscle, and a well-tufted oil-gland. This specimen was kindly given me by Canon Tristram, carbolized and dry. Of genera which have already passed through my hands I have had the following additional species :— Ara militaris. Licmetis tenutrostris. Cacatua moluccensis. Lorius domicella. philippinarum. Paocephalus meyert. Hclectus grandis. Tanygnathus albirostris. Eos reticulata. They all agree with those species previously dissected, except Licmetis tenwirostris, which has only one carotid, the left, whereas L. pastinator has two. It will be interesting to verify this difference between the South-Australian species and its more western ally ; for the uncertainty of the disposition of these vessels in the Cacatuine is rendered more striking if it is correct. In the specimen dissected, of Cacatua philippinarum, a gall-bladder was present. Thisis the only case in which I have seen this viscus in any Parrot. Formulating the varying characters of the above newly dissected genera upon the principle adopted in my earlier paper and there ex- plained, the formule run thus :-— (1) Dasyptilus 2.—.+.+. (4) Chalcopsitta 2.—.+.+. (2) Deroptyus 2.—.+.+. (5) Coriphilus 2.—.+.+. (3) Polyteles 2.—.+.+. Such being the case, Dasyptilus and Deroptyus fall into my sub- family Pyrrhurine, whilst the other three must be placed with the Palwornithine. It is interesting to notice that Deroptyus agrees with Pyrrhura, and not with Conurus. EEE ON THE GALL-BLADDER OF PARROTS. 263 41. NOTE ON THE ABSENCE OR PRESENCE OF A GALL-BLADDER IN THE FAMILY OF THE PAR- ROTS.* In a former communicationt I had the opportunity of showing that Page 793. the generalization, founded upon the dissection of an insufficient number of genera, that the gall-bladder is wanting in the Columba, does not apply to Carpophaga, Lopholemus, or Ptilopus. On the present occasion I have to correct a similar error with reference to the Psittaci, because I have found a well-developed gall-bladder in speci- mens of Cacatua-philippinarum, Cacatua goffini, Cacatua moluccensis, and Calopsitta nove-hollandie, in which last-named species it is small and easily overlooked. _ In my earlier dissections I have not recorded the presence of a gall-bladder in any species of Parrot. That, no doubt, is because, it being absent in so many, I did not expect to find it. From the above facts it is highly probable that the presence of a gall-bladder in the Cacatuine will have to be included among the _ characteristic peculiarities of this subfamily. At the same time its persistence in them is in favour of the view that the Palzornithinz, as restricted by me,{ are but little different from the ancestral Parrots, and the Cacatuinz still less so. The primitive Parrots must have possessed a gall-bladder—because we know that this organ is present in the Cacatuinz, and consequently was not absent in the primitive species, as the probability that it should have been independently developed a second time is infinitely little. * “Proceedings of the Zoological Society,” 1877, p. 793. Read, Nov. 20, 1877. + “ Proceedings of the Zoological Society,” p. 257. (Supra, p. 239.) t “ Proceedings of the Zoological Society,” p. 594. (Supra, p. 255.) 264 ON HALMATURUS LUCTUOSUS. 42, ON THE KANGAROO CALLED HALMATURUS LUCTUOSUS BY D’ALBERTIS, AND ITS AFFI- NITIES.* (Plates VIII-X.) | Page 48. Durine the time that H.M.S. “ Basilisk” was cruising in the region of the south-east of New Guinea one of the sailors acquired a speci- men of a small Kangaroo, which Signor L. M. D’Albertis, C.M.Z.S., obtained from him at Sydney. In a letter addressed to Mr. Sclater, dated Sydney, N.S.W., December 1, 1873, Signor D’Albertis de- scribed this specimen, under the name of Halmaturus luctuosus, as followst :—“ Length from the nose to the occiput 43 inches; length of the ears 13 inch; length of the thigh 52 inches; length of the tarsus, including the nail, 43 inches; length of the tail 114 inches. Total length, from the nose to the tip of the tail, 2 feet 5 inches. Its weight is 74 pounds. “The fur is short, its general colour dark ashy brown with a silvery tinge, white at the roots; chin, throat, and chest white, with two horizontal ashy stripes under the pouch; on the top of the head a silvery whitish spot; the thighs more grey; feet dark, almost black; the arm white inside; the hand black. The tail moderately strong, of a similar colour to the body, but white and bare of hairs for about an inch at the extremity. The lips are barely covered with fur; the eyelids are puffed, almost naked, and provided with eyelashes so fine as not to be readily seen at first sight.” Hab. “8.E. of New Guinea.” On April 17, 1874, this Kangaroo was deposited by Signor D’Al- bertis in the Society’s Gardens; and at the Meeting for Scientific Business on May 5th following, Mr. Sclater, in reporting on the additions to the Society’s Menagerie, exhibited a drawing of it, and referred to it as the “typical example of Halmaturus luctuosus of D’Al- bertis.” It is this specimen, a female, which forms the subject of the present communication. It died, Nov. 24, 1874, with congested lungs, after a severe frost, the first of the commencing winter. An examination of the dead body, and especially of the mouth, which it was impossible to observe in the living animal, made it * “Proceedings of the Zoological Society,” 1875, pp. 48-59. Pls. VII—IX. Reed, Feb. 2, 1875. + “ Proceedings of the Zoological Society,” 1874, p. 110. t “ Proceedings of the Zoological Society,” 1874, p, 247, Pl, XUIL. EEE EE ——s ON HALMATURUS LUCTUOSUS. 265 evident that the species could not be rightly included in the genus Macropus or Halmaturus. Further comparison made it clear that it was intimately related to the genus Dendrolagus, and also to the species described in Waterhouse’s “‘ Mammalia ”* as Macropus brumii. Pege 49. Mr. Waterhouse bases his description of this last-named species on a skin so labelled in the British Museum, and on Miiller’s account of the same animal in his elaborate work,t in the letterpress of which it is termed Dorcopsis brunii. The priority of the generic name being undisputed, any fresh species which can be shown to be generically related to the above-determined species is evidently a species of the genus Dorcopsis. This last remark is called for because the subject is rendered somewhat involved by an oversight of the illustrious Miller. In his description of his Dorcopsis brunii he evidently has no doubt that the Specimen or specimens he is considering, is or are identical with the “Philander” described by Bruynf as having been seen by him in the garden of the Governor of Batavia, upon which the name brunii was originally based. Prof. Schlegel,§ however, has most convincingly shown the unjustifiableness of this assumption, and has proved beyond a doubt that the species to which the name Philander can alone be applied is that found only in the islands of Aru and the Ké group, _ whilst the species which forms the subject of Miiller’s memoir is a ~ denizen of New Guinea itself. Prof. Schlegel therefore retains the name Macropus brunii for the Philander of Aru, and of the New- Guinea animal forms the new species Macropus muelleri. As to me it is evident that M. muelleri is generically distinct from Macropus in its widest sense, and from all its minor divisions, it is also evident that Dorcopsis muelleri must be the name applied to the Dorcopsis brunii of Miller. The species which forms the subject of the present com- munication, belonging (as I hope to prove) to the same genus as Dorcopsis muelleri (Schlegel), must therefore stand as Dorcopsis luctuosa (D’ Albertis). The material at my disposal is the following:—the skin and skeleton of the type specimen of Dorcopsis luctuosa; the skins of an adult male and female, as well as of a young male, of Vorcopsis muel- lert in the British Museum, collected by Mr. Wallace; a skull from the skin of the above-mentioned female of Dorcopsis muelleri; the much-discoloured skin of the male of the same species in the British Museum, from New Guinea, described by Mr. Waterhouse|] as Macro- * Vol i. “ Marsupiata,” p. 180. + “ Zoogdieren van den Indischen Archipel,” pt. 4, Pl. XXT. } “ Reizen over Moskovie,” p. 374, Pl. 213 (1713). § “ Nederlandsch Tijdschrift voor de Dierkunde,” 1866, p. 350 ef seq. || “ Mammalia,” vol. i., p. 180. Page 50. 266 ON HALMATURUS LUCTUOSUS. pus bruni; two skeletons of Dendrolagus inustus, one in the British Museum and the other in the Museum of the College of Surgeons; as well as a pair of skins and an imperfect skull of Macropus brunit from Aru, kindly lent me by Mr. Edward Gerrard. So faras I know, the visceral anatomy of Dorcopsis muelleri has not been described. That of Dendrolagus inustus is fully given by Prof. Owen in the “ Proceedings” of the Society ;* and some of the actual specimens on which this description is based are preserved in the Museum of the College of Surgeons. The internal anatomy of Macropus brunii is not known. The following Table gives the most important measurements of the skin of the female Dorcopsis luctuosa, compared with specimens of the same sex of Dorcopsis muelleri and Macropus brunii :— Dorcopsis | Dorcopsis | Macropus Lengths, ‘Eo. luctuosa 3. | muelleri 9.| brunii 9. in. in. in. From tip of nose to base of tuil..........006. 24 20°25 21:0 Daa hike Tae se sei SaGialeie ea elceies Reman 13 25 15 °4 11°75 From tip of nose to occiput... 1... ..eeee eens 5:0 5°0 4°0 BOre GIGS. 5 oss 40s pss nt aoeunineiatacente 5°75 6°75 4°75 Hind limb i uate oo ak Shas BA he aka ome 10°75 12°55 10°5 From heel to end of nail of fourth toe........ 4°75 4°75 5°0 LeeniBth OF OB8, aic.0sc sok ph onan Sie motrin 1°4 1°25 1°75 Circumference of base of tail ............... 4°25 ae 2°0 From knee to knee over the back............ 14:0 17°0 The general contour of the body is quite Macropine; the breadth at the hips, however, is somewhat small. The hair is soft, short, and of a nearly uniform length all over the skin. The head is elongate and conical, the muffle naked, the eyes large and antilopine. The colour of the upper surface and sides of the headt and back is uniformly blackish with a silvery gloss, each hair being whitish at its base for two fifths of its length, black for the next two fifths, and white at the tip. On the ventral surface a broad longitudinal white band extends from the line joining the angles of the mouth, backwards along the neck and belly as far as the pouch, behind and from the sides of which it continues towards the tail of a true slate-colour as far as the cloacal orifice, between which spot and * “Proceedings of the Zoological Society,” 1852, p. 103 et seq. + The silvery white spot on the top of the head, mentioned in D’Albertis’ description, is not produced by the presence of white hair, but results from the fact that the spot where it is sometimes seen is the anterior junction of the forward- directed hair of the neck with the backward-directed hair of the frontalregion. Its existence depends entirely. on the way in which the hair is brushed; and it is not visible except after the natural disposition has been disturbed. ON HALMATURUS LUCTUOSUS. 267 the base of the tail it is again white. This white band occupies the whole of the region between the angles of the jaw, and continues down the neck over the abdomen of a slightly greater width. It only encroaches on the sides of the body by sending an expansion into each axilla, which is visible laterally just behind the elbow. There is no lateral transverse white stripe across the front of the thigh, like that so strongly marked in M. brunii; and, unlike this last named species, the light grey, nearly white stripe above and parallel to the lip is very insignificant, and does not extend backwards under the eye. The ear is rounded, black inside and ont, with a slight white line _ formed by the similarly coloured roots of the there exposed hairs bounding the auditory meatus anteriorly. - The non-exposed surfaces of both the arm proper and the thigh : are of a pale grey. The other parts of both the fore and hind limbs are black. The nails of both the fore and hind limbs are short and Macropine. The peculiarity in the direction of the hair of the neck, which Page 51. elsewhere occurs only in Dorcopsis muelleri, Dendrolagus ursinus, and Dendrolagus inustus, is as strongly marked as in those species—all the hair covering the space bounded in front by a line running trans- versely across the parietal region, and behind by two lines joining in the middle line between the shoulders to form a right angle seven ~ inches behind the occiput, and extending forward and outward to the shoulder-joint, being directed forward, whilst the general body-cover- ing of hair is directed normally backwards. The lips are nearly naked, as is the skin covering the subsym- j physial portion of the mandible, just behind which are four large ’ and conspicuous glandular hair-follicles in the middle line, arranged in pairs to form a square (Plate [9] VIII.). A collection of glands of a similar nature is found on the upper eyelid, situated a little nearer : the inner than the outer canthus. These are shown in Miiller’s drawing of Dorcopsis muelleri.* A few long hairs are to be found on the sides of the upper lip. The eyelids are somewhat puffed, almost naked, with the eye- lashes scarcely apparent. The tail is peculiar in being of considerable diameter to near its extremity, and in being uniformly thickly covered, for all but its termination, with soft, not very short, black hair. The skin of the distal end of the tail is black, except for its terminal 14 inch, where it is nearly white. On the upper part of this white portion there are a few white hairs; elsewhere it is naked and scaly. The scales are also distinctly seen extending forward for a short space over the * Loc. cit. Pl. XXII. Page 52. 268 ON HALMATURUS LUCTUOSUS. inferior surface of the black skin, from the absence of hair in that part. The characteristic manner in which the animal employs its tail as a method of support (well shown in “ Proceedings of the Zoological Society,” 1874, pl. xlii.), might have almost been predicted from the above-described distribution of the hair; for it is evident that only a part at the extreme end could have habitually come into contact with the ground. The only brown hair on the body is that in the pouch, which is rufous. There are four mamme. There is not the least difficulty in distinguishing Dorcopsis luctuosa from D. muelleri. The general colour of the head, back, and tail in the specimens of the latter species from Mysol, above referred to, is a mouse-chocolate, which becomes duller over the thighs, and of a pale grey on the outside of the fore limb. In D. muelleri the general white of the abdominal surface expands slightly opposite the orifice of the pouch, just above the knees; it, however, does not develop into a band over the flank as in Macropus brunii: the white of the throat also extends on to the angle of the jaw, and continues forward to join the dim white stripe along the upper lip; and there is a second insignificant white line under each eye, also (as mentioned by Prof. Schlegel) not nearly so marked as in M. bruni. In the male of D. muellert the white tip to the tail is as much as three inches in length. The skull of Dorcopsis luctuosa (Plate [8] VII.) very closely resembles that of D. muellert, the following being the two most im- portant measurements in adult specimens of the same sex (female) :— D. luctuosa. D. muelleri. in, in. Length of eleull. . .s «ise ssw sioee 41 4°55 Greatest breadth, from zygoma to CY GOLGR ns 55 Ais mis ea eit a cop panels 2:2 2°05 In some minor details there are slight differences. In D. muelleri, as in most species of Macropus, the premaxillary region is bent down- wards in such a way that the line formed by the trenchant edges of the molar teeth, if projected onwards to the nose, is quite above the incisor teeth. In D. luctuosa this bending downwards of the snout is not so marked, as will be seen by comparing the side view of the skull (Plate [8] VII. fig. 3) and the similar one of D. muelleri in Prof. Miiller’s elaborate work above referred to. The palatine foramina, one large one on each side, together with several much smaller ones behind each, in D. muelleri end behind the transverse palato-maxillary sutures, whilst in D. luctuosa their anterior margins are formed by the palatine plates of the maxillary bones, into which they encroach a short distance. In D. luctuosa the upper LS — ON HALMATURUS LUCTUOSUS. 269 of the lacrymal foramina in each lacrymal bone has an ossific ridge behind it, which causes it to be completely exserted, or situated on the face outside the orbit; whilst in D. muelleri the absence of this bony ridge causes it to be situated in a recess on the margin of the orbit. In D. luctuosa the apex of the angular process which is developed downwards from the inferior margin of the maxillary portion of the zygoma, is opposite the anterior cusp of the third molar tooth, whilst in D. muelleri it corresponds to the posterior cusp of the second molar. With regard to the teeth themselves, the canines in D. muelleri are quite the size of or even slightly larger than the most lateral incisor ; in D. luctwosa, however, they are much smaller, being nothing more than slightly curved dentine cylinders about 3; of an inch in diameter, as in the subgenus Lagorchestes, and directed downwards and forwards. In both the species the third incisor has an inflection on its labial surface, as in all the species of Macropus: in D. muelleri this fold is a little in front of the middle of the tooth ; and in D. luctuosa it is decidedly nearer the posterior border. In the last-named species there is a simi- lar distinct inflection on the second incisor; in D. muelleri this is not apparent. In D. muelleri the inferior incisor is directed more imme- diately forward than in D. luctuosa, in which it turns slightly upwards; this peculiarity is correlated with the difference in the obliquity-of the " premaxillary region (vide Plate [10] IX.). In the enormous premolars there is a slight difference—those of D. muelleri being a little the larger, in the upper jaw having a breadth of 0°55 inch against 0°475 inch for the same teeth in D. luctuosa. In D. muelleri the bony septum between the two fangs of each premolar, especially of the lower jaw, is particularly conspicuous in the undis- Page 53. turbed tooth, even projecting slightly beyond the osseous alveolar margin. In D. luctuosa this septum is scarcely visible. The most important characters of the skull of Dorcopsis, as a genus, which distinguishes it from Dendrolagus, are the following :— In Dendrolagus the head is proportionally much shorter, the effect of which on the lower jaw is that, as the dental series is not correspond- ingly reduced, the ramus and the body of each lateral moiety meet at a right instead of an obtuse angle; there are no palatine foramina; the zygoma is considerably deeper; the exoccipital processes are longer, though not much so; the lower incisors are considerably broader, at the same time that the upper lateral incisors are larger and more cylindrical, with superficial grooves which can scarcely be termed inflections; the premolars are not so broad, and their outer posterior tubercles are more distinctly developed. The molar teeth of Dorcopsis and Dendrolagus are almost identical (vide Plate [10] TX.). Page 54. 270 ON HALMATURUS LUCTUOSUS. The cranial characters which distinguish Dorcopsis, as a genus, from Macropus, are not very significant. Looking at the base of the skull the arrangement of the teeth deserves attention. In Dorcopsis the premolar with the molars on both sides form straight lines, which are exactly parallel one to the other; whilst in Macropus the molar- premolar series form slight curves, convex outwards, converging behind as well as in front.. In Dorcopsis the zygomata are not so powerful or deep from above downwards, as in the similar-sized species of Macropus. A peculiarity also presents itself in the lateral occipital region, the exoccipitals descending considerably below the free extremities of the paramastoids in Macropus, whilst in Dorcopsis they reach downwards scarcely any further distance. : Respecting the teeth, Dorcopsis differs from Macropus in the much diminished size of the superior lateral incisors. The central incisors are not so broad, but nearly as long. The second incisor is very much smaller ; and though presenting a slight inflection in D. luctuosa, as mentioned above, this inflection is not, as in Macropus, posterior and internal, at the line of contact with the anterior margin of its more lateral neighbour. The third incisor is also very much smaller. The inflection on its labial or outer surface presents the same differences in . the two species of Dorcopsis that are found in the various species of Macropus: in D. luctuosa, as in M. brunii and M. thetidis, it is very near its posterior border; whilst in D. muelleri, as in M. major and most of the other species, it is far forward. The inferior incisors in Dorcopsis are proportionally narrower than in Macropus, in which peculiarity Dendrolagus resembles the latter genus: they, however, wear down in a similar manner, namely at the anterior end of the supero-lateral margin, differently from that in the Hypsiprymniform Macropodide, in which they wear in a rodent-like fashion. The presence of the superior canines in Dorcopsis distinguishes it from most of the species of Macropus, although they are almost as well developed in the subgenus Lagorchestes as in D. luctuosa, and in that one only. The premolars of Dorcopsis are particularly interesting, presenting characteristic features which are more suggestive of its affinities than any other skeletal point. As to those in the upper jaw, their breadth from before backwards is very nearly or quite as great as that of the first and second molar, together with the anterior of the two cusps of the third. The crown of the tooth on each side is prismatic in shape, with one of the angles forming the cutting-edge, the oppo- site side the base. A tubercle on the inner surface of the posterior end of the tooth disturbs the uniformity of the prismatic shape; it is ON HALMATURUS LUCTUOSUS. 271 continued forward along the margin of the lingual surface as a feebly developed ridge or cingulum. Opposite it on the labial surface, a small tubercle is also to be found, larger in Dendrolagus, with a similar, slighter cingular expansion. From the thus somewhat swollen neck or cingulum several ridges with intervening depressions run at right angles, to end at the trenchant edge. These ridges differ considerably from those observed in the corresponding tooth of the genus Bettongia, in other points than their degree of obliquity: they are less numerous, and therefore further apart, because the tooth is considerably broader; and they are continued as what look like tumefactions of their basal ends, into both the inner and outer cingulum. It may be here mentioned that the premolars of Hypsiprymnus proper (H. murinus, H. gilberti, and H. platyops) agree much more closely with Dorcopsis and Dendrolagus in the characters in which those genera differ from Bettongia. The mandibular premolars are much like those in the maxilla. They are not so broad, equalling only the two succeeding molars. They present a tumefaction or cingulum at the base of the crown; but the posterior internal and external tubercles are not developed. In Macropus there is never anything like the size of the premolars of Dorcopsis or Dendrolagus, although there is a considerable range of _ difference in different sections of the genus, which in Macropus proper ~ appears to me to be correlated with the length of ear rather than with any other character. In M. major their size and permanency is but slight; and in most of the long-eared species they are not so broad as the first true molar. In M. billardieri and M. brunii they attain their maximum size proportionally; and they are nearly as large in the subgenera Petrogale and Lagorchestes. In M. billardieri and M. brunii they are almost exact miniatures of those in Dorcopsis, except that the number of perpendicular ridges is fewer. Respecting the molars of Dorcopsis and Dendrolagus, they may be termed macropodiform, because, though much resembling those of the type genus, they present special characters. The two transverse pris- | matic ridges, with the small connecting bridge between them, are ; present, although the last-named structure is less conspicuous and narrower. The anterior minor ridge is also to be seen ; itis, however, Page 55. much smaller and narrower than in Macropus; as in that genus, it is more marked in the mandibular than in the maxillary molars. The peculiar twist in the molar-premolar series of the lower jaw (the ante- rior teeth turning outwards and the posterior inwards), by which the trenchant edges are rendered parallel, as in the upper jaw, at the same time that the rami of the mandible converge is, as might be expected from the previously mentioned greater parallelism in the maxillary series of Dorcopsis, more marked in that genus than in ee 272 ON HALMATURUS LUCTUOSUS. Macropus. It may be mentioned that the molar teeth in Dorcopsis and. Dendrolagus do not exhibit any characters intermediate between Macropus and Hypsiprymnus. ' The remaining bones of the skeleton do not present features of special interest. The typical number of precandal vertebree are pre- sent, namely, C. 7, D. 13, L. 6, and S. 2; there are 19 candal vertebra, with well developed chevron bones between the proximal ones. The anterior arch of the atlas presents no gap, the two moieties meeting with a linear junction. The anticlinal vertebre are the 10th, 11th, and 12th dorsal. The clavicles are fairly developed; and the first ribs are very broad. There is asupracondyloid foramen to the humerus ; and the fibula is not ossified to the tibia. The following are the lengths of some of the most important long bones :—Humerus, 2°75 inches, radius 3°2, femur 5:1, tibia 6-2, fourth metatarsal 1:8, pubic symphysis 1°7. Respecting the soft parts, the tongue has three small circumyallate papille at its base, arranged in the ordinary V-shaped manner. The palate presents several strongly marked transverse ridges. The sub- maxillary and sublingual glands are small, the former ellipsoid in shape. The parotids are large, flat, and triangular, with their bases directed towards the root of the neck, and their apices to the masseter muscle. Their position is indicated by the dotted lines in Plate [9] VIII. The left lung is formed of a single lobe, with a slight fissure on the ' ventral margin, near the apex, opposite the broadest part of the heart. Page 56. The right lung consists of two lobes, the main portion and the azygos lobe. The lobe proper presents two fissures—one near the apex, running vertebrally and diaphragmatically, separating an apical lobule, the other running vertebro-apically, and marking off the median lobule.* This median lobule partly embraces the base of the heart, as in many animals. There is no third bronchus. The heart is quite Macropine, there being two superior vens cave. The right ventricle also spirally wraps round the much stronger left, as in Macropus. The stomach is perfectly Macropine; that is, it is elongated, sac- culated, with the cesophagus entering it munch nearer the cecal than the pyloric extremity, with the walls of the pyloric end smooth and much thickened. The cardiac cecal extremity, like that in Dendrolagus as described by Prof. Owen, consists of a single cul-de-sac, not a bifid one like that in Macropus giganteus. * The method of description here adopted is an attempt to avoid the employ- ment of terms, which necessitate any assumption with respect to the position of the animal. Supposing the animal to have its vertebral column horizontal, and its four limbs on the ground, then the above description might be thus read :—“ The lobe proper presents two fissures—one near the apex, running upwards and backwards, the other running forwards and upwards.” ora rer ees ON HALMATURUS LUCTUOSUS. 273 In the subgenus Petrogale the stomach is not bifid at its cardiac extremity, in which respect it resembles Dorcopsis. In other respects, however, it presents considerable differences; it is more capacious opposite the cesophageal orifice, and the cardiac portion is bent on the rest nearly at right angles, which is not the case in Macropus giganteus and Dorcopsis. The character of the mucous membrane also deserves attention.* In Macropus giganteus, as is well known, the squamous epithelium of the cesophagus spreads over most of the stomach also, the pyloric extremity, and one of the two cardiac ceca (which is itself bifid) being alone lined with a columnar coating. In Petrogale this latter is absent, the digestive mucous membrane being confined to the pyloric region. Of Dendrolagus inustus Prof. Owen remarks,t “the epithe- lium is continued from the cesophagus, for a breadth of 2 inches down the posterior surface of the stomach, and of 1} inches down the ante- rior surface, and thence is continued, slightly diminished in breadth, 3 inches towards the pyloric end of the stomach, and 23 inches towards the cardiacend. The rest of the cavity is lined with the usual gastric vascular membrane, the surface of which is diversified by patches of follicular apertures along the upper curvature of the stomach, which patches increase in breadth as they approach the true digestive portion.” A very similar condition maintains in Dorcopsis luctuosa, the only difference being that the squamous lining covering the whole of the cardiac cul-de-sac is also found to spread from the esophageal orifice along the lesser curvature for a short distance towards the pylorus. As in Dendrolagus inustus, two strong parallel longitudinal folds run from the cesophageal opening, in this squamous- covered mucous membrane, for some distance on the way to the pyloric compartment, gradually disappearing before they reach it. The small intestine is 97 inches in length, with numerous oblong Peyer’s patches distributed throughout its whole distance, averaging 1} inch long, by } inch across. The cecum and large intestine are not sacculated ; the former has a length of 24 inches, and its cireum- ference is the same; the latter is 32 inches long, being one-third the length of the small intestine, which is the same proportion that Prof. Owen{ observed between the same-named viscera of Dendro- lagus inustus. The equally short cexcum in the Hypsiprymni differs in having two lateral longitudinal bands which scarcely sacculate it. * [See, for a full description of this, the paper by Messrs. E. A. Schafer and D. J. Williams “ On the Structure of the Mucous Membrane of the Stomach in the Kangaroos,” which was communicated to the Zoological Society by Prof. Garrod, “* Proceedings of the Zoological Society,” 1876, pp. 165-177.— Eb. ]} + “ Proceedings of the Zoological Society,” 1852, p. 105. t Loe. cit. p. 106. Page 57. 274 ON HALMATURUS LUCTUOSUS. The spleen is perfectly Macropine, being narrow and elongate, with a well-developed third lobule. The liver very closely resembles that of the different species of Macropus. In comparing the livers of different animals it is my habit to estimate by sight, and therefore only approximately, the bulk of the different lobes, and to write down the results in the form of a formula. Employing the divisions, so evidently natural, proposed by Prof. Flower, I commence by writing down the name of the largest lobe, after which the others in the order of their bulk, with symbols between each to indicate their relative size. Taking the liver-formula of Dorcopsis luctuosa as an example, it may be thus written, L.L. 2>C. 4>R.C. $>Sp.>R.L. 2>L.C. ; and it reads as follows :—The left lateral lobe (.L.) is the largest ; it is twice the size of the caudate (C.), which is half as large again as the right central (R.C.), which is half as large again as the Spigelian(Sp.), which is larger (very little) than the right lateral (R.L.), which is twice the size of the left central (L.C.). The similarly constructed formula of Macropus melanops is L.L.=C. 2>R.C. $>R.L. 3 >Sp. 3>L.C., and of Halmaturus derbianus L.L. 1>C=R.C. $>Sp. $>R.L. 2>L.C.: they show how great a similarity there is between the different mem- bers of the family Macropodide. The gall-bladder is situated in the deep cystic fossa; and the um- bilical fissure is not deep. The Spigelian lobe has its apex directed vertebrally and resting on the left lateral lobe, as in Macropus; no secondary lobules are connected with it. There is a peculiarity in the liver of the specimen of Dorcopsis luctuosa under consideration, which may be individual, or it may be characteristic of the species, genus, or subfamily; at all events, I have not seen it in any other mammalian animal. Looking at the diaphrag- matic surface of any multilobate liver, the lateral margins of the mass formed by the right and left central lobes are always seen to overlap, to a greater or less extent, the lateral lobes in an imbricate manner. Similarly the right lateral lobe overlaps or covers the caudate. In the livers of Macropus and Halmaturus which I have by me, this conforma- tion is strictly maintained. But in Dorcopsis luctuosa the caudate lobe overlaps the right lateral lobe (instead of being situated on its abdominal surface), in such a way that the last-named lobe is only seen between the right free edge of the right central lobe and the left free edge of the caudate. This condition is not brought about by any post mortem change in the position of the lobes, because the right lateral fissure is not so deep as to separate them at their vertebral extremity. ON HALMATURUS LUCTUOSUS. 275 The uterus is perfectly Macropine, as are the vagine. No direct communication could be found between the uterine pouch of the vagine and the common vaginal canal. A gland, as usual, about the size of an almond, with a slender duct, opens on each side of the narrow cavity included between the sphincter ani and the external common sphincter. — Tn conclusion, the comparison of the various organs and structures of the Macropodide which have come before me in my study of Page 58. Dorcopsis luctuosa would lead me to divide up the family in the fol- lowing manner :— Family MACROPODID. Diprotodont Marsupialia wanting the hallux, the second and third digits of the pes being much reduced and included in the skin as far as the ungual phalanges, which at the same time have the claws so formed that the inner is convex inwards and the outer convex outwards, at the same time that their contiguous surfaces are flattened. The stomach is elongated and sacculated. Subfamily Macropop1nz. Macropodidz in which the cesophagus enters the stomach near the cardiac end; with a Spigelian lobe to the liver; with no lateral longitudinal bands to the colic cecum when it is short, and with radius of normal form. Section 1. Macropus. With the premolars never much larger than the first molar; with a characteristic molar-tooth-pattern ; with the stomach but slightly lined with digestive epithe- lium (?) and with the hair on the nape of the neck directed backwards. Hab. Australia, Tasmania, Aru, and the Ké Islands. Genera or subgenera. Macropus, Halmaturus, Petrogale, Lagor- chestes. Section 2. Dorcopsis. With the premolars strikingly large, with a characteristic molar-tooth-pattern, slightly modified upon that of Macropus ; with the stomach mostly lined with diges- tive epithelium, and with the hair of the nape of the neck directed forwards. Hab. New Guinea and Mysol. Genus 1. Dorcopsis. Limbs Macropine in their proportions. Genus 2. Dendrolagus, Protemnodon*, Sthenurus*. Fore limbs much longer than in Macropus. * An inspection of the plates in Prof. Owen’s paper on these new genera (“ Phil. Trans.,” 1873, p. 245), makes it evident that they are scarcely distinguish- able from Dendrolagus, and must be included in the Doxcopsis section of the family. Tt 2 Page 59. 276 ON HALMATURUS LUCTUOSUS. Subfamily HypsipryMNIN&. Macropodide in which the ceso- phagus enters the stomach near the pyloric end; with no special Spigelian lobe to the liver; with lateral longitudinal bands to the short colic cecum; with a much-flattened and expanded radius, with a characteristie molar-tooth-pattern, and with the incisors worn down much as in Rodent animals. Hab. Australia and Tasmania. Genus Hypsiprymnus (including H. murinus, H. gilberti, and H., platyops). Auditory bulla somewhat inflated ; palatine foramina, one large one on each side; ridges on premolars few and perpendicular. Face elongate. Genus Bettongia (including all the others of the group except B. rufescens). Auditory bulla much inflated; palatine foramina as in Hypsiprymwus; ridges on premolars numerous and oblique; head short. Genus MWpyprymnus* (including only Bettongia rufescens of Gould). Auditory bulla net inflated; palatine foramina absent; head short; tarsus considerably longer than in the two other genera. It should be mentioned that the viseeral anatomy of Apyprymnus rufescens has not been published, and that Mr. Waterhouse divides the genus Hypsiprymnus into three subgenera corresponding exactly with the three genera here defined. My best thanks are due both to Mr. Sclater and Dr. Giinther for the very kind way in which both these gentlemen have assisted me in my study of this subject. EXPLANATION OF THE PLATES. Prats 8. (VII) Lateral superior and inferior views of the skull of Dorcopsis luctuosa, natural size. Prate 9. (VIII.) View of the inferior surface of the neck of Dorcopsis luctuosa, showing the median gland with four orifices situated in the hyoid region. The positions of the large parotid and small submaxillary glands are indicated by dotted lines. PuatEe 10. (TX.) Teeth, twice the natural size, of (figs. 1-5) Dorcopsis luctuosa, (figs. 6-10) Dorcopsis miilleri, and (figs. 11-15) Macropus brunii. The upper two rows represent the left upper premolar, the third and fourth rows the upper and lower third left molar, and the bottom row the incisors. * This term I propose for Mr. Waterhouse’s first section of Hypsiprymnus, which he has ieft without any Latin name. ON THE MECHANISM OF THE BIRD’S WING. 277 43. ON A POINT IN THE MECHANISM OF THE BIRD'S WING.* Te beautiful investigations of Borelli, together with those of Page 82. - M. Marey, make it certain that in any organ which is employed as a flapping wing there must be a stiff or rigid anterior margin. In the insect the stout anterior nervure performs this function; in the bird the bones of the arm, forearm, and manus do the same. How, in the latter, this necessary rigidity is developed, considering the presence of the elbow- and wrist-joints, must be, at first sight, a matter of surprise. It depends on the mechanical arrangement by which, when, in the wing, the arm is bent on the forearm, the manus is always similarly bent on the forearm; and when extension of the forearm is made, extension of the manus equally certainly follows. This occurs when all the muscles and tendons are removed, and the ligaments binding the bones together are alone left. The explanation of this mechanism is not difficult. The arm con- sists of one bone only, the humerus; the forearm of two, the ulna and radius ; the manus of the two carpals together with the metacarpals and phalanges. The mutual relations of these two bones are such that the radius and ulna move one above another like the two limbs of a pair of drawing-parallels, each being fixed proximally to the humerus and distally to the carpus. The plane common to the radius and ulna is the same as that in which flexion and extension of the elbow is per- formed, so that one of the two bones of the forearm, the radius, articulates with the humerus at a point nearer the shoulder, or further ~ from the elbow, than its companion, the ulna. At the wrist the radius . Is consequently superior, articulating with the carpal bone on the pollex side: whilst the ulna articulates with the other element of the carpus. This condition maintaining the parallel movements of the radius on the ulna must necessarily be attended by a parallel move- ment of the humerus on the manus. When the humerus bends upon the ulna, the manus therefore similarly bends upon the forearm; and the triceps muscle is able, unassisted, to maintain the whole limb in a rigid state during extension. In making a wooden model of these bones to illustrate the above described mechanism, one or two points of mechanical detail suggested a reference to the shape of the distal end of the humerus. The wing = « Proceedings of the Zoological Society,” 1875, pp. 82, 3. Read, Feb. 16, 1875. 1 278 ON THE MECHANISM OF THE BIRD’S WING. Page 83. in the living bird, when at rest, is completely folded; and when fully extended forms but a slightly angular rod. To allow of this consider- able range of movement of the bones of the forearm on the humerus, and of their being completely folded up, it is necessary to attach a very projecting hinge at the portion of the model of the humerus which represents the humero-ulnar articulation, otherwise, when fully flexed, the model radius would not be able to be included between the then parallel humerus and ulna; especially as the radius, to get in its fully flexed position, must rotate on a hinge which itself projects its semi- diameter at least beyond the humerus. These requirements explain the characteristic shape of the distal end of the humerus in birds. It is curved towards its flexor side, and sharply so at its extremity where it comes in contact with the ulna. At the same time the radius: articulates with it on a well-developed knob, situated above the similar surface for the ulna, and to its outer side (which allows of a less extensive joint). The similar arrangement required at the wrist-joint is arrived at by the inter- polation of the carpal bones between the forearm and consolidated metacarpus. In some wings, when all the muscles are removed, this movement is not so manifest as in others, there being a certain amount of in- dependent power of movement in the manus in all positions. This is much reduced in the living bird by the tendon of the tensor patagit . longus muscle, which runs from the shoulder, along the free margin of the patagium, to the wrist, where, in being attached to the metacarpal mass on the pollex side, it aids the extension of the manus during the extension of the forearm. The mechanism above deseribed is stated by Dr. Alix* to have been first indicated by Bergmann, as far as the anatomical arrange- ment is concerned, although Strauss-Diirckheim, in his ‘Théologie de la Nature’ was the first to explain it fully. Dr. Alix himself has alsot entered into the detail of the movement “of elongation ” of the radius, which is well explained in his large work above referred to.t My object in bringing the subject before the Society is to draw special attention to so important a point, and to illustrate its action by a wooden model, which demonstrates its accuracy in a very striking manner, It may be here mentioned that the movement of the general plane of the wing during both the up and down stroke, which by Borelli and his followers is ascribed to the elastic yielding of the feathers in birds, * ‘ Essai sur l'appareil locomoteur des Oiseaux,” Paris, 1874, p. 230. + “ Bulletin de la Société Philomathique,” 1864. t Loe. cit. p. 330 et seq. ON THE TRACHEA OF CERTAIN DUCKS. 279 and of the wing-membrane in insects, appears to me rather to be de- pendent on the torsion of the bones or main nervure of the wing, the power of lateral flexion in which is proved by M. Marey’s discovery of the figure-of-8 action in the insect. A thin wooden lath employed as a nervure to an artificial wing, if set with its narrow section vertical and fixed to a non-yielding horizontal wing, gives a vertical figure- of-8 when moved up and down, the plane changing exactly as it is described by M. Marey in the insect. 44, ON THE FORM OF THE LOWER LARYNX IN CERTAIN SPECIES OF DUCKS.* THE present communication contains descriptions of the condition of Page 151. the lower larynx in some rare members of the Anatide, which are not referred to in the works of either Mr. Eyton or Mr. Yarrell. 1. Sancrp10Ryis MELANONOTA (Gm.): Sclater, Rev. Cat. Vert. p. 241. Page 152. To Mr. Eyton, who established the genus to which this peculiar bird _ belongs, the visceral anatomy was unknown; and I am not aware of _ any subsequent description of it having been published. A pair were purchased by the Society on the 18th of September, 1867, the female of which died on the 10th of March, and the male on the 18th of October last year; these are the specimens which I have examined. In both sexes the diameter of the trachea diminishes slightly at its lower extremity before it again expands a little to end in the syrinx. As in birds generally, the tracheal rings are complete and notched in the middle line before and behind, in such a way that where they meet the two halves overlap and are overlapped respectively by the rings above and below them. The lower tracheal rings, however, in both sexes are much thinned in front, as is the case in the male of Harelda glacialis ;+ they are not ossified together. In the male Sarcidiornis melanonota (fig. 1) there are 20 anterior, membrane-covered fenestre, formed in the intervals between these thinned rings; in the female (fig. 2) there are only 12 of the same. Page 153. In the latter there is no lateral diverticulum from the syrinx; but in the former, from the left side, as usual, one is developed, entirely a we * “ Proceedings of the Zoological Society,’ 1875, pp. 151-6. Read, March 2, 1875. + Vide figs. Eyton’s “ Anatide,” plate opposite p. 65; Yarrell’s “ Brit. Birds,” vol. iii. p. 261. 280 ON THE TRACHEA OF CERTAIN DUCKS. osseous, irregularly compressed, and very small, not having a di- ameter in any part greater than that of the trachea itself (vide figs. 1 and 2). _ Fig. 2. Fig. 3. a oe. > et beonlll Femmes = al ar Pi: Dl Dagegt = (gent sd ‘T) a re TELS i ee. Mipediddtidisite mene = Fig. 1. Lower part of trachea of Sarcidiornis melanonota @ . Fig. 2. Ditto of Sarcidiornis melanonota ¢ . Fig. 3. Ditto of Rhodonessa caryophyllacea $. In the male specimen the ceca are 3 and 2} inches long; in the female not quite 2 inches. Their diameter is inconsiderable, not exceeding 1 of an inch. The whole intestinal canal measures between 44 and 5 feet; and the gizzard is decidedly small, not being bigger than that of a common Duck (Anas boscas). 2. RHODONESSA CARYOPHYLLACEA (Lath.). Anas caryophyllacea, Scl. ‘‘ Proceedings of the Zoological Society,” 1874, p. 110. This rare Duck is generally placed in the genus Anas; by Mr. Eyton, however, it is considered to belong to the Fuliguline ; and that ornithologist puts it, along with Fuligula rufina, in the genus Callichen. A pair purchased by the Society on the 12th January last year, died, the female on the 11th and the male on the 15th of March, 1874. From these two specimens I was able to remove the windpipes for examination. The structure of the syrinx of the male is in favour of == th Mi 5, ON THE TRACHEA OF CERTAIN DUCKS. 231 the Faliguline affinities of the genus; and the trachea presents points Page 154, of superficial similarity to that of the last-described bird, Sarcidiornis melanonota, as will be seen by a comparison of the accompanying draw- ings (figs. 1-5) of the lower portion of the windpipes in the two. In the female (fig. 3) there is no lateral diverticulum, the syrinx being Fig. 4. Fig. 5. HL com en een pany ae eo a por eal ; iin “er oo —_— — =. — — ional — es —— i b id aageepiye’ Mba eatiy Fig. 4. Lower part of trachea of Rhodonessa caryophyllacea of (front view). Fig. 5. Ditto (side view). simple. The lower end of the trachea is hardly contracted at all. There is, however, aslight thinning of the anterior portions of some of the inferior tracheal rings, as in the female of Sarcidiornis melanonota, though to a less extent—a small, transverse, anterior fenestra being the result. In the Rhodonessa the syrinx proper is nevertheless differently constructed, the last five or six tracheal-rings being con- solidated together, the fenestration being situated higher up; whilst in the Sarcidiornis the fenestration of the unanchylosed rings con- tinues as low down as the bronchial bifurcation (vide figs. 3, 4, and 5).. In the male Rhodonessa caryophyllacea (figs. 4 and 5) the lower portion of the trachea is less capacious than a little higher up, where a slight fusiform dilatation occurs. Above the large syringeal box there are in front 15 transverse fenestre formed between the thinned tracheal rings, as in the Sarcidiornis and Harelda. Below them the syrinx is formed by a considerable dilatation in two directions—one to Mpyes « ON THE TRACHEA OF CERTAIN DUCKS. 282 Fig. 6. Page 155. ii heA HNN) i wananed rf ny \ i We) Tue Ge | 7 Sh A, mi AeA Trachea of Metopiana,peposaca of (front view). ee 5 (Sere ON THE TRACHEA OF CERTAIN DUCKS. 283 the left, which is the larger and has semimembranous walls; the other slightly to the right, inferior in position to theformer. This latter is simply osseous, no fenestree being present in it; it intrudes upon the right: side as well as the left in front. The last 12 orso tracheal rings are considerably dilated and co-ossified, the two above-mentioned compartments being connected with the cavity formed by their fusion through a single left-sided orifice, the left bronchus springing from the membraniform cavity. The cxca are not quite equal in size, being 23 and 1? inches long; the whole intestine measured 4 feet. 3. Meroprana Peposaca (Vieill.): Sclater, Rev. Cat. Vert. p. 255. Of this bird Mr. Sclater mentions* that “it has a large bulbous expansion in the windpipe.” This I have found in all the male specimens which I have examined. Its distance above the bifurcation of the bronchi is best estimated from the accompanying sketch (fig. 6, p- 282) which is of the natural size. A similar tracheal dilatation is to be observed in the male of Melanitta fusca, that in Clangula his- trionica being much less considerable. In a male, purchased on the 6th of July, 1870, which died on the 7th of January last, the syringeal Fig. 7. a lel Lower part of trachea of Metopiana peposaca ¢ (side view). * “ Proceedings of the Zoological Society,” 1868, p. 146. Page 156. Page 297. 284 ON THE TRACHEA box (see figs. 6 and 7) is constructed on the same type as in Fuligula rufma and fF. ferina, being mostly composed of membrane, with an intersecting, oblique, simple osseous bar running across near the upper margin of its outer side. There is also some dilatation of the consolidated rings which go to form the lower portion of the trachea; this is to be observed on both the right and left sides, the box being connected with the latter only. In the female no box is developed. The trachea narrows slightly above the syringeal box. The cwca in this specimen were 5} and 6 inches in length, the whole intestinal canal measuring 44 feet. 45, ON THE FORM OF THE TRACHEA IN CERTAIN | SPECIES OF STORKS AND SPOONBILLS.* No account of the peculiarities of the windpipe in Tantalus ibis and in Platalea ajaja has yet, to the best of my knowledge, appeared in print. They cannot but interest ornithologists; I therefore append descriptions of them from specimens which have passed through my hands as Prosector to the Society. In the Transactions of the Linnean Societyt there is a paper by Mr. Joshua Brookes, F.R.S., ‘On the remarkable Formation of the Trachea in the Egyptian Tantalus.” The author does not mention the sex of his specimen, and does not refer to the existence of any intrathoracic or any other loops; he draws attention only to the existence of a lateral compression of the portion of the trachea which is contained within the thorax; and he incidentally refers to the similarity of the arrangement of the windpipe in the Spoonbill and Tantalus ibis, but does not hint at the points in which they agree. In most species of Ciconiide the only peculiarity of the windpipe is that the bronchi are longer than in other birds, the bifurcation of the trachea occurring at, or even a little above, the superior aperture of the thorax. This condition I have observed in the female Oiconia boyciana which died on January 15th, 1874, as well as in examples of C. maguarit and C. alba. In the male of C. nigra the bronchi are known to be peculiarly long,t and to form an m-shaped curve enter- * “ Proceedings of the Zoological Society,” 1875, pp. 297-3801. Read, April 6, 1875. + Vol. xvi. p. 499. t{ “ Naumann’s Naturgeschichte der Vogel Deutschlands,” vol. ix. p. 229. IN STORKS AND SPOONBILLS. 285 ing the lungs. No other peculiarities have been described among these birds. : A specimen of Tantalus ibis was purchased by the Society on the 26th of May, 1873, which died on the 12th of March, 1875. It proved to be a male. The following is the arrangement of the convo- Intions of its trachea (see figure, p. 286). The windpipe descends the neck in front of the cesophagus without any peculiarities being observable, the rings which go to compose it being exactly like those of other allied birds, circular, complete, elastic, notched in the middle line before and behind, and of ordinary depth. Directly it reaches the superior aperture of the thorax, between the two rami of the furcula, a sudden change occurs. The succeeding rings are inelastic, from being ossified; and they are ossified together in pairs, so that their apparent depth is more than double that of the cervical rings, the intermediate membrane being included in the double rings. The depth of the unmodified rings is hardly more than ~, of an inch, that of the intrathoracic modified ones being as much as } of an inch. The diameter of both is about 4 of an inch; those in the chest are further peculiar in developing a slight median longitudinal ridge along their posterior surface. The two musculi depressores trachece, after ranning down the wind- Page 299. pipe as long as it is in the neck, leave it together as it enters the chest to run to their insertions behind the sternal articular ends of the second complete ribs, the left one crossing in front of the upper of the two loops described below. There are no special lateral muscles running to the syrinx. The trachea, modified in the manner above described, continues its normal course downwards as far as a horizontal plane cutting the base of the heart, when it makes a fairly gradual turn through half a circle, directly forward, to consequently ascend with the posterior keel above mentioned, running along the middle of its convex surface. On reaching the level of the symphysis furcule it makes a second semi- circular turn to the right, to again descend nearly as far as_on the former occasion, and making a third similar turn to the left whilst in the fold of the first loop, ascends a third time as high as the line joining the two sides of the furcula—in other words, to the very top of the thoracic cavity. Here it turns backwards to descend again, in contact with the first part of the intrathoracic tube, to its right side, as far as the level of the apex of the heart and the com- mencement of the proventriculus ; where, making a short very abrupt turn forwards, it bifurcates into the two bronchi, which therefore, uniquely, run from their origins upwards and outwards to their re- spective lungs. In this third and last descending portion of the windpipe, which has a length of 53 inches, the lower 3 inches are — i al 286 ON THE TRACHEA Page 298. f Fig. 1. wt 4 ae Intrathoracic convolutions of the trachea in Tantalus ibis. e, coracoid; f, furcula; p, proventriculus; 7.b, right bronchus; /.4 left bronchus; sf, sternum. IN STORKS AND SPOONBILLS. 287 considerably flattened in what would be the lateral direction, which through the convolutions it has been called upon to make, is twisted, ~so that the flattening appears to be nearly antero-posterior, the median ridge, developed posteriorly, being placed considerably on the right side. : About an inch above this flexure, in which the bronchi bifurcate, the previously deep double rings suddenly cease to be developed as such, and return to their normal condition just before the peculiarly situated and simple syrinx is reached. There are altogether 82 of the ossified double rings in the modified portion of the windpipe. The earlier bronchial rings are peculiar in being deep, the fibro- cartilaginous rings being ossified and thickened above and below for a certain portion (the external) of their circumference. Platalea ajaja.—The peculiar convolution, within the thorax, of the trachea in Platalea leucorodia is well figured by Mr. Yarrell.* The arrangement in Platalea ajaja is, however, quite different. A pair of these birds was purchased by the Society on the 13th of August, 1870. The female dying on the 27th of July, and the male on the 13th of October, 1873, have given me the opportunity of ex- amining the windpipe in both sexes. The trachea is simple, straight, of uniform calibre, and peculiarly short, extending only two-thirds down the length of the neck, where the uncomplicated syrinx is situated and the bifurcation of the bronchi occurs. The usual pair of muscles, one on each side, runs to this syrinx from above, and ceases there. The bronchi are fusiformly dilated at their commence- ment, where the rings which encircle them are not complete, a mem- Page 301. brane taking their place in that portion of each tube which is con- tiguous to its opposite neighbour. Each bronchus, lower down, is composed of complete cartilaginous rings (vide fig. 2, p. 288). By many ornithologists Tantalus is arranged along with Platalea and Ibis, instead of with the Storks. Nitzsch, in his “ Pterylo- graphy,” places it with Oiconia in his group Pevarci, separating off Platalea and Ibis to form the Hemigtormmes. In the “ Revised List” of the Animals in the Society’s Gardens, Mr. Sclater adopts the same arrangement. In my paper ‘“‘On the Nasal Bones of Birds,” f it is mentioned that Platalea and Ibis are schizorhinal—that is, have the external osseous nares split up in a manner there described, in which point they differ from the rest of Prof. Huxley’s Pelargomorphe, and therefore from Tantalus. There are many other structural peculiarities which make it per- * “ British Birds,” vol. ii. p. 504. + “ Proceedings of the Zoological Society,”’ 1873, p.33. (Supra, p. 124.) 288 ON THE TRACHEA IN STORKS AND SPOONBILLS. Page 300. Fig. 2. —b Cervical bifurcation of the bronchi in Platalea ajaja. a, trachea; 6, syrinx; d, cesophagus; e, cervical muscles and vertebre ; r.b, right bronchus; 7.6, left bronchus. fectly certain that Tantalus is a member of the Ciconiide, and not an aberrant one either. Some of the most important it will not be out of place to mention here. They will be most easily appreciated in a tabular form, as thus represented :— ON THE DEEP PLANTAR TENDONS IN BIRDS. 289 is thee end Pistalon. The skull is schizorhinal. The angle of the mandible is produced and recurved. The pectoralis major muscle is simple, not being separable into distinct layers. The accessory femoro-caudal muscle is well developed. | The semitendinosus muscle is muscular throughout. A small muscular belly is sent from the biceps cubiti to the In Oiconia and Tantalus. The skull is holorhinal. The angle of the mandible is truncated. The pectoralis major muscle is in two layers, a superficial and a deep, easily separable one from the other. The accessory femoro-caudal muscle is absent. The semitendinosus muscle is tendinons for its distal half. No slip leaves the biceps cubiti muscle to join the tensor patagii tendon of the tensor patagii longus longus. muscle. 46. ON THE DISPOSITION OF THE DEEP PLANTAR TENDONS IN DIFFERENT BIRDS.* ~ ‘Te arrangement of the tendons in the palm of the hand and the Page 339, sole of the foot among the Mammalia is a subject of great intrieacy, as may be inferred from the comparison of the dissections of different Page 340. animals whose anatomy has been sufficiently investigated. Among birds peculiarities in the disposition of the plantar tendons has already attracted the attention of Professor C. J. Sundevall, who, as is well known, divides the Passeres off from all other orders, and includes Upupa with them, because in them, and in them only, the tendon of the flezor longus hallucis musele is quite independent of that of the flexor perforans digitorum ; whilst in other birds the former joins,the latter, so preventing the two from being quite independent in their 2 action. Ali other descriptions which I have seen of special dissections > have been confirmatory of this view; and my own observations, with but a slight exception in the case of Botaurus, to be mentioned below, support Professor Sundevall’s separation off of the Passeres together with Upupa on this particular character. My dissections, however, have shown me that there is still more to be learnt from the plantar tendons, and that the large mass of birds which all agree in that the two above-mentioned deep flexors blend together, present among * “ Proceedings of the Zoological Society,” 1875, pp. 339-48. Read, April 20, 1875, U he Page 341, 290 ON THE DEEP PLANTAR TENDONS IN BIRDS. themselves peculiarities as important as that which so definitely characterizes the Passeres. To describe and to endeavour to show the bearing of these differences are the objects of the present paper. In birds generally, whatever the number of their toes, there are two muscles whose fleshy bellies are situated in the leg proper (that is, between the knee and the ankle), deep, and just behind the tibia. These muscles arise, one from almost the whole of the posterior sur- face of the tibia and from the fibula, in a bipenniform manner, and the other from the inferior surface of the horizontal femur, just be- hind the outer genual articular condyle. The former is termed the flexor perforans digitorum pedis, because its terminal tendons per- forate those of the more superficial flexors on their way to the ungual phalanges of their respective toes; and the latter is termed the flewor longus hallucis, because there is generally a shorter muscle to the same digit. These two muscles descend to the ankle (the joint between the tibio-tarsus and the tarso-metatarsus) side by side; they run behind it, in the fibro-cartilaginous or osseous mass which, in birds, is always found at the posterior part of the upper end of the tarso-metatarse, in two canals, deeper than any of the other flexor tendons; and in these canals there is always a definite relation between them. Some- times the tendons are side by side; and then it is always that of the flewor longus hallucis which is the external of the two, the osseous vertical ridge, which is nearly always seen in the dry bone, separating them. Sometimes, however, one is superficial or, in other words, posterior to the other. When this is the case it is always the flexor perforans digitorum which is the deeper. In the Swifts, for instance, the flecor longus hallucis quite covers the flexor perforans digitorum ; but in most Parrots, as may be seen by the disposition of the osseous canals in the dry tarso-metatarse, that for the former muscle is exter- nal as well as superficial, only partially covering it. These relations are constant, and must be always borne in mind in all attempts to identify the muscles. From these it can be inferred, as is verified by dissection, that the tendon of the flewor longus hallucis crosses its companion superficially on its way from the ankle to its insertion in the hallux. Just before, or just at the commencement of, the sole of the bird’s foot (near the joint between the metatarsus and the phalanges), these two tendons generally split up to supply the toes. By far the majority of the families of birds agree in the distribution of the terminal tendons, conforming to one common type. This typical arrangement must be first described. The common Fowl (Gallus bankiva) is a very good example. The accompanying diagram (fig. 1) will assist in explaining it. The tendon of the flezor longus hallucis descending on ON THE DEEP PLANTAR “TENDONS IN BIRDS. 291 the outer side of the tendon of the flexor perforans digitorum, crosses it Fig. 1. Gallus bankiva. superficially in its downward and inward course to the lower surface of the base of the hallux, whence it traverses the flexor surface of that digit to the base of the ungual phalanx at which spot it is inserted. The flexor perforans digitorum continues down to the sole of the foot as a single tendon, where it immediately splits into three parts, one to the ungual phalanx of each of the three anteriorly directed digits. Opposite the lower part of the tarso-metatarse the Page 342. flexor longus hallucis sends downwards a fibrous vinculum (V) which joins the flexor perforans digitorum tendon just before it commences to trifurcate. In all cases this vinculum is always directed downwards from the hallux-muscle to the digits-muscle, so that, when the tendon of the flexor perforans digitorum alone is pulled upon, the three anterior digits alone are flexed; but when the flexor longus hallucis is put in action, the digits as well as the hallux are simultaneously flexed. The proportion borne by this vinculum to the main tendon of the - flexor longus hallucis varies considerably. In some birds it is com- paratively feeble and insignificant ; whilst in others, with but a small hallux, it is much larger than the hallucial moiety, and seems to be u 2 Page 343. 292 ON THE DEEP PLANTAR TENDONS IN BIRDS. the main continuation outwards of the insertion of the muscle into that of the flezor perforans digitorum, the slip to the great toe being but small compared with it. In the Dorking Fowl the flexor longus Fig. 3. Fig. 4. il Tinnunculus alaudarius. Buceros rhinoceros. hallucis tendon splits into two (after it has given off the vinculum to the flexor perforans), one resulting portion going to the normal hallux and the other to the supplementary toe, which is therefore a hallux also, as is generally supposed. This manner of distribution of the deep plantar tendons, which is that found in a great number of birds, may be summarized as follows :—The flexor perforans digitorum splits opposite the meta- tarso-phalangeal joint into three tendons, one running to the ungual phalanx of each of the three anteriorly directed toes. The flexor longus hallucis is inserted into the ungual phalanx of the hallux, but it sends downwards near the middle of the tarso-metatarsus a vinculum to join the tendon of the flexor perforans digitorum just before the tri- furcation of that muscle (figs. 1 and 7). This condition is fonnd in the following birds which I have examined :— | Gallus bankiva. Musophaga violacea. Megacephalon maleo. Schizorhis africana. Fulica atra. Orotophaga sulcirostris. ON THE DEEP PLANTAR TENDONS IN BIRDS. 293 Pheenicophaés, sp. ? Leptoptilus argala. Eudynamis orientalis. Ardea sumatrana — very ~ Cuculus canorus. slender). Nestor notabilis. "cinerea (vinculum scarcely Chrysotis festiva. exists). , ochrocephala. Cancroma cochlearia. Ara chloroptera. Geopelia cuneata. Baza lophotes. Ibis rubra. Syrnium aluco (vinculum ‘very Platalea ajaja. broad). Eurypyge helias. - In Ardea cinerea and in A. sumatrana, here mentioned, the vinculum is stated to be extremely feeble. In Botawrus stellaris this condition is carried a step further, the vinculum being quite wanting. Prof. Sundevall states that such is the case only in the Passeres and in Upupa ; here, however, is a slight exception to that generalization. Frequently the vinculum above referred to is so considerable in strength that it makes the flezor longus hallucis appear to fuse with the flexor perforans digitorum, and only to send a slip before doing so to the hallux. This condition is evidently but an inconsiderable modi- fication upon the previously described typical arrangement (fig. 2, p- 291). It is, however, a stepping-stone to others, which it assists in explaining. It is found in the following birds, which I have dissected (it will be noticed that they have the hallux comparatively a a cant) :— Palani mantelli. Cygnus nigricollis. & Nothura maculosa. fF odiceps minor. = Chenalopez egyptiacus. Phalacrocoraz carbo. In many of the Accipitres Diurne a slight modification of this arrangement is observed. The flezor longus hallucis divides into two moieties opposite the lower end of the tarso-metatarse, one of which runs to the hallux. The other partis the representative of the vinculum of the above-mentioned birds; it is peculiar, however, in that, instead of joining the tendon of the flexor perforans digitorum before it is dis- tributed to the anterior toes, it mostly runs down to blend with the slip which is associated with the inner of these (digit 2) only (fig. 3). This condition I have observed in Haliaétus albicilla, Tinnunculus alaudarius. In Geranoaétus aguia and in Polyborus brasiliensis, besides the special tendon from the hallux-muscle to the second digit, there is a Page 344. broad thin vinculum present, as in Gallus. In the Accipitres Diurne the arrangement of the tendons therefore differs in different groups— 294 ON THE DEEP PLANTAR TENDONS IN BIRDS. in Baza their distribution being quite normal, that is as in the first- described manner ; in Polyborus, Haliaétus, Tinnunculus, and Geranoétus this condition is combined with a special extra tendon to the second digit, which greatly increases its power of flexion. The arrangement observed in the Cathartide is in no way allied to any of these, and adds another important point to the many now known to separate them off entirely from the Accipitres vere. The next arrangement to be described is a very different one. The two deep flexors descend beyond the ankle-joint independently, as usual; after passing which, generally about one-third down the tarso- metatarse, they blend completely, before any slip has been given off. From the conjoined tendon thus formed the tendons of distribution spring, four in number, one to the hallux and others to each of the three anteriorly directed toes (fig. 4, p. 292), that to the former being generally separated off before any of the others. Among Homalogonatous birds the only group in which I have observed this condition is that of the Cathartide—both Cathartes atratus and Sarcorhamphus gryphus possessing it, and so differing entirely from their supposed allies the diurnal Accipitres. Among Anomalogonatous birds the arrangement is very commonly found; I have seen it in Coracias garrula, Podargus cuviert, Buceros rhinoceros, Caprimulqus europeus, Steatornis caripensis, Cypselus alpinus. On looking at the plantar tendons thus arranged, without further dissection, the slip to the hallux from the conjoined deep flexor tendon seems to spring from its inner (that is, hallucial) side) ; whereas, from what has been said above, the long flexor of the hallux is situated external to the common flexor, at the ankle-joint. Further, in these birds, on straining upon the distal hallux slip with one hand, at the same time that the distal slips to the remaining toes are held in the other, the two elements of the conjoined tendons: tend to divide up in the direction of the ultimate fibres ; and in doing so the line of rupture always develops in such a way that it leaves the thus further-separated hallux slip still on the inner side in connexion with the main flexor perforans tendon. . A natural condition, like’ this thus artificially produced one, is found in some birds closely allied to those in which the last described arrangement obtains. It is found in Momotus lessoni, Dacelo gigantea, and Merops apiaster. In them the tendons of the flexor longus hallucis and of the fleeor perforans digitorum pass down beyond the ankle-joint in the typical manner, the former external to the latter as usual. Opposite the upper end of the tarso-metatarse the flexor perforans digi- ON THE DEEP PLANTAR TENDONS, IN BIRDS. 295 torum gives off from its inner side the flexor slip which supplies the hallux, the majority of the tendon descending as usual towards the foot. Opposite the middle of the tarso-metatarse it is joined by the Page 345. tendon of the flexor longus hallucis on its outer side, whereupon the : conjoined tendon splits into three divisions to supply the three anterior toes (vide fig. 5). The peculiar conformation in the foot of the Trogonide is associated with an equally abnormal arrangement of the plantar tendons, which I have found in Trogon puella and in Pharomacrus mocinno. In these birds the tendon of the flexor longus hallucis is situated, as it ought to Fig. 5. Fig. 6. Iv ut IV Il Momotus lessoni. Trogon puella. be, external to the flexor perforans digitorum ; it also crosses it super- - ficially, opposite about the middle of the tarso-metatarse, sending down a slender vinculum in the normal manner. The peculiarity is in the ultimate destination of the tendons, the flexor longus hallucis and the flexor perforans digitorwm each dividing into two near the metatarso- phalangeal articulation, the two portions of the former tendon running to the hallux and digit 2, the two of the latter to digits 3 and 4 (vide fig.6). This arrangement is not found in any other group of birds, as far as my experience goes. 296 ON THE DEEP PLANTAR TENDONS IN BIRDS. Besides the three last peculiar arrangements of the tendons, which T have not found elsewhere described, there is another still more pecu- liar and unexpected. I have observed it in all the Anomalogonatous Page 346. birds with scansorial feet which I have examined, and in them only, it being present in Ramphastos ariel, Tiga javanensis, Megalema asiatica, Galbula albirostris, Gecinus viridis, Urogalba paradisea. It is represented in fig. 8. The two tendons descend behind the ankle as usual, having their origins typical. There is nothing peculiar till they have descended two-thirds down the tarso-metatarse. About opposite the middle of that bone the flewor longus hallucis sends a vinculum downwards as in the Fowl, to join the tendon of the flexor perforans digitorum. Just above the metatarso-phalangeal articulation the tendons become arranged for distribution in a most uncommon manner., The tendon of the flexor perforans digitorwm does not split up, but runs to one digit only, namely the third toe, which is the Fig. 7. Fig. 8. Til Crotophaga sulcirostris. Megalema asiatica. outer of the two that are directed forward. It is covered superficially by the flecor perforans digitorum, just as that latter muscle is splitting up to be distributed to the hallux as well as to digits 2 and4, In these ON THE DEEP PLANTAR TENDONS IN BIRDS. 297 birds we have, therefore, the flexor longus hallucis arising from the lower surface of the femur only, running through the ankle at the outer side of the other deep tendon, and sending a vinculam down- wards—all of which are special characters of that muscle only, it being Page 347. distributed to three toes, whilst the flexor perforans digitorum only supplies one. The birds with scansorial feet thus fall into two divisions, accord- ing to the arrangement of their plantar tendons, these being normal in the Psittaci and Cuculide, whilst they are extremely peculiar in the Picide, Ramphastide, Capitonide, and Galbulide. In my paper on the Classification of Birds,* the presence or absence of the ambiens muscle made me feel justified in placing the Psittacit and Cuculide among my HomaLoconar2, at the same time that the Pici, Ramphastide, Capitonide, and Galbulide are arranged among the ANOMALOGONATZ. These new observations are therefore strongly in favour of the natural- ness of the classification proposed. There is only one other point to be considered on the present occa- sion, as far as this question is concerned. It is the distribution of Fig. 9. a pill A typical Passerine Foot. * “ Proceedings of the Zoological Society,” 1873, p. 626, and 1874, p. 111 e¢ seq. (Supra, p. 187, et seq.) Page 348. 298 ON THE DEEP PILANTAR TENDONS IN BIRDS. these tendons in birds which do not possess the hallux, or in which there is no long flexor tendon to that digit when it is present. In all these cases both the flezor longus hallucis and the flexor perforans digi- torum muscles are present and well developed, only they blend com- pletely opposite the upper part of the tarso-metatarse to form a single common tendon to be distributed, on its splitting up, to the anterior toes—to the two of Struthio, the three of Rhea, Otis, &c. Dr. Alix* has described how that, in the common Swan (Cygnus olor), there is no long flexor tendon to the small hallux. I have not examined that species; but there is undoubtedly a small one in OC. nigricollis, C. atra- tus, and in all the other Anserine birds I have examined, as above mentioned. However I have found this tendon to the hallux wanting in Parra africana, Pygosceles papua, Chauna derbiana, Podiceps minor. Professor C. Sundevall has shownt that in the Passeres and in Upupa epops the tendons of the flexor longus hallucis and the flexor perforans digitorum are quite free from one another, not being united by any vinculum. In all the Passeres which I have examined my observations agree with these generalizations. However, the same condition maintains in Botawrus stellaris and almost in Ardea cinerea, where the vinculum is scarcely more than a single fibre (vide fig. 9). DESCRIPTION OF THE FIGURES. In all the figures the numbering refers to the digits I, II, III, IV, representing the hallux, second, third, and fourth digits respectively. In all, the deep plantar tendons are alone represented, and these from their plantar aspect, the hallucial tendon being the outer of the two at the heel-joint. Fig. 1. Left foot of Gallus bankiva; V, vinculum running downwards from the outer hallucial tendon to the inner digital common tendon. . Right foot of Apteryx mantelli. . Right foot of Zinnunculus alaudarius. . Right foot of Buceros rhinoceros. Right foot of Momotus lessoni. Arrangement of the tendons in the left foot of Trogon puella. . Right foot of Crotophaga sulcirostris. . Right foot of Megalema asiatica. . Right foot of a Passerine bird. CONA KE ww * < Rssai sur l’appareil locomoteur des Oiseux.” Paris, 1874, p. 464. + “ Methodi naturalis avium disponendarum tentamen” (Stockholm, 1872), and elsewhere. ON THE HYOID BONE OF THE ELEPHANT. 299 47. ON THE HYOID BONE OF THE ELEPHANT.* Tae hyoid apparatus of the Indian Elephant (Hlephas indicus) Page 365. present peculiarities which a study of the same in the Ungulata would tend to complicate rather than to simplify. The basihyal together with the thyrohyals form an arch (of which, by the way, I have not seen the components anchylosed even in adult specimens)—which does not present the least difficulty, a small pair of cartilaginous lesser cornua being present in the position of the lesser cornua of anthro- potomy. It is the stylohyals which, as far as I can find, have not yet been correctly described. Of them Professor Owen remarksf, “‘ From the middle of the stylohyal a slender pointed process is sent off at an Page 366. acute angle.” And in Professor Flower’s “ Osteology ” it is said that “the stylohyals are remarkable for having a long pointed process projecting downwards from near the middle of their posterior border.” Professor Morrison Watson{ enters fully into the de- scription of the hyoid muscles, without mentioning, though evidently correctly understanding, the disposition of the bones with which they - are associated. From the above remarks it is evident that the thick short portion of each stylohyal is assumed to be the body of that bone, the pointed process being considered to be an accessory part of it. Such, how- ever, is not the case—the slender pointed process in reality corre- sponding to the long body of the stylohyal in the Perrisso- and Artiodactyla, whilst the short thick process is the posterior descending process. That such is the case I have been able to prove recently in two specimens. In the Indian Elephant the stylohyal (s h) is a thin bone composed, Page 367. in the adult male, of a flat portion, 4 inches long, less than } inch thick, and very nearly 2? inch broad, with parallel sides, obtusely truncated at both ends, which are capped with cartilage. In the standing animal the position of this portion of the bone is nearly ver- tical. Above, it is closely united to the stylo-temporal region of the skull, whilst the lower end gives origin to the digastric muscle. From the middle of the anterior border continues onwards the body of the bone, at an acute angle with the lower portion of the above-described * “ Proceedings of the Zoological Society,” 1875, pp. 365-7. Read, May 4, 1875. + “ Anatomy of Vertebrates,” vol. ii. p 441. - t “ Journal of Anatomy and Physiology,’ November 1874, p. 131. 300 ON THE HYOID BONE OF THE ELEPHANT. element, downwards and forwards. This is elongately triangular in shape, 6 inches long, $ inch broad at its middle, and tapering to a Hyoid bones in Indian Elephant. point in front, where it gives attachment to a hardly specialized stylo- hyoid ligament and serves for the origin of the stylo-glossus muscle. The interval between the tip of this stylohyal and the lesser cornu (cartilaginous) of the hyoid bone is 5 inches, or a little less than the length of the process itself. As it descends in its downward and forward course, this tapering stylohyal curves slightly on itself, turn- ing a little outwards. The accompanying figure will explain the condition. The descending digastric process, as it may be termed, may be compared to the posteriorly directed process of the stylohyal in the Ungulata. It differs from it, however, in one essential particular, which is that in the latter it does not give origin to the digastric muscle, but only to the stylohyoid; whilst in the Elephant the digas- REPORT ON THE INDIAN ELEPHANT. 301 tric arises from its lower end only, and the stylohyoid from the angle formed at its junction with the body of the bone. In the Elephant therefore the deficiency of the lateral interme- diate elements of the hyoid apparatus permit of a much greater movement of the base of the tongue than in the Ungulata, whose nearly rigid stylohyals, epihyals, and ceratohyals can allow of little more than an antero-posterior movement of the base of the tongue, in part of the circle of which the hyo-cranial attachment is the centre. 48. REPORT ON THE INDIAN ELEPHANT WHICH DIED IN THE GARDENS JULY 71a, 1875.* On May Ist, 1851, the Society purchased of Mr. Batty (then of the Page 542. Circus, Westminster Bridge), for £800, an adult female Hlephas indicus with its female calf. The specimens had been deposited in the Society’s Gardens on the 19th of the preceding month. In the spring of the year 1850, John Stimpson, now keeper in the Society’s service, left the E.I. Company’s military service, and when at Cawn- poor, on his way to Calcutta, met an animal-dealer, Mr. Wallace, who was on his way to Calcutta with the female and calf in question as well as another Elephant. Stimpson is sure that the calf was born after the female had been captured, and thinks that it was three : months old when he first saw it. He assisted in taking charge of the animals till they arrived in this country: they were five months on the voyage. ee Of the two specimens purchased by the Society the mother was sold on April 28th, 1854, to the Zoological Society of Brussels, the calf continuing to suckle until that date, i.e., until upwards of four years of age. It is this calf of 1851 which died on July 7th, 1875 (25 years old). The Superintendent, the head keeper, and the Elephant-keepers are of opinion that it continued to grow until within a year of its death. Its height at the withers at the time of its death was just eight feet. For the last four years at least the animal has lost the power of extending its trunk, from paralysis of the anterior intrinsic muscles of that organ. It has thus not been able to throw its trunk over its * “Proceedings of the Zoolegical Society,” 1875, pp. 542-38. Read, Nov. 2, 1875. Page 543. 302 REPORT ON THE INDIAN ELEPHANT. head, or even the least forwards. When it took food it flexed the trunk so as to present the orifice forwards. This symptom is one of decay. For the last two years of its life it exhibited marked signs of rheumatism, varying in severity, very considerable at times. This was most manifest on its kneeling down to be saddled. “pin The animal during the last three years of its life looked preter- naturally aged and worn out. It has never suffered the least from cough, and has not become strikingly thin. It carried its saddle and visitors 36 hours before its death, apparently without discomfort, and ate well on the evening of the 6th inst. For about six months the animal did not, as it was formerly wont to do, lie down at night. On the night of the 6th of July it fell on its left side, and did not subsequently make any powerful attempt to rise. The breathing was, when down, unusually rapid (about 25 a minute); and no marked symptoms of pain manifested themselves, general discomfort being evident. It died during the night of the 7th, having at 8.30 p.m. had a large dose (over 100 grs.) of strychnia given it by the mouth. Whether the poison was the cause of death is uncertain. 3 With the exception of one of the lungs, all the organs, the brain included, were perfectly healthy. The lung in question was almost entirely infiltrated with tubercular deposit, not more than one-sixth being competent for the respiratory function. The tubercular infil- tration was uniform or nearly so, being of a lighter colour and nearly in a condition to break up in the centre of the organ, forming a dark grey solid mass in the more recently affected portions near the margins. It may be mentioned that the teeth just coming into wear had, in both jaws, 23, 24, or 25 plates; they were therefore the sixth molars. The epiphyses of the long bones were firmly united. No entozoa were found. As to the duration of the disease it is not easy to decide. It, no doubt, was of considerable standing ; probably it had commenced with the first signs of decrepitude, about three years ago. As to the cause of the tuberculosis, that was probably connected with the animal having been born and bred in captivity in a cold climate. A wild-caught animal 4 or 5 years of age would probably have thriven better. ON THE MANATEE. 303 49. NOTES ON THE MANATEE (MANATUS AMERI- CANUS) RECENTLY LIVING IN THE SOCIETY’S GARDENS.* (Plates XI—XIIL.) Unt: the arrival, on August the 6th of this year, ofa living female Page 137. Manatee in the Society’s Gardens, very few Europeans had had the opportunity of seeing a live Sirenian; and those who had previously been so fortunate, had not,in most cases, been able to observe it under the favourable circumstances of position afforded by a small shallow pond approachable on all sides. In this pond it was not the least difficult to watch the creature minutely whilst it was feeding, as well as to make other notes of its habits which would necessarily be over- looked by those who meet it in its native haunts. Dr. Maurie, in his valuable monograph on the Manatee,t justly remarks, that “in museums it is customary to see such bloated over- stuffed specimens, that from them, as well as figures extant, an unfair idea of the configuration is obtained.” Dr. Murie’s own illustrations, with their beantiful textural details, represent the animal much more accurately. The only fault in them, if it is a fault, is rather in the opposite direction to that above indicated by him. His specimens were dead and preserved; they had evidently shrunk slightly, the living animal appearing a little more rounded, and presenting less well- marked skin-folds. The following are the measurements of the animal which I have had the opportunity of dissecting, taken almost immediately after death :— Potal lengths Sen AA MK HRS: 80 Circumference (3 inches in front of umbilicus) 50 Greatest breadth of tail (8 inches from end).. 19 Umbilicus to anus ......... aes sBWene ek baled 21 Walvai to ema tetas. Fit. Bee eas eee ss 4°25 Circumference at root of tail (7 inches beyond SHGB). (3. Pa; See Pet Se PRLS 28°75 Length of tail-flapper: ..........0....5...00. 19°5 Length: of fore Timbo... S20. Soe ie. res ok 14 From eye to front of snout ................ 4°75 * “Transactions of the Zoological Society,” X. pp. 137-145. pls. XXVIII— XXX. Read, Nov. 16, 1875. + “ Transactions of the Zoological Society,” VIII. p. 127. Page 138. 304 ON THE MANATEE. Tnches Breadth of snout halfway down ............ 55 Height of snout at middle line.............. 3 From ear to ear over head ..............0. 8°75 From eye to eye above head................ 6°4 From ear to eye on one side................ 5°75 Whether in Dr. Murie’s figured specimen the snout and front part of the head had been swollen during life, whether the enlargement was the result of post mortem change, or whether it was on account of the youth of the individual, that part of the body is represented of considerably larger proportionate size with reference to the rest of the animal than it was in the Society’s living example, in which the head . was more distinctly like a clean-cut truncated cone, without any well- marked transverse folds or appearance of puffiness. It is, however, in the oral margin of the upper lip that the dead- ness of Dr. Murie’s specimen is. most manifest; and the fact that the peculiarity in the mechanism of that organ has not yet (so far as I am aware) been described, indicates how impossible it sometimes is to predict special functions from anatomical structure alone. The upper lip is prehensile; in other words, the animal is able, by its unaided means, to introduce food placed before it into the mouth without the assistance of the comparatively insignificant lower lip. To understand how this is accomplished it is necessary that the struc- ture of the labial margin should be described. A front view of the head, Plate [11] XXVIII, fig. 2, shows that on each side of the narrow median portion of the superior labial margin of the oral orifice depends a rounded lateral lip-pad of con- siderable size, which gives a deeply notched appearance to the upper lip. Of these lip-pads, Dr. Murie tells us* that “at the dependent angle on each side of the muzzie is a circumscribed oval prominence half an ineh in diameter, where the ridges, furrows, and bristles [found elsewhere] are specially pronounced. This spot would seem to possess most tactile delicacy; for twigs of the infraorbital and facial nerves are abundant thereto. . . . . The above dispo- sition strongly reminds one of the moustachial apparatus of the Walrus; but their shortness and rigidity render them unequal to perform the office of a sieve, as is the case in the Pinniped: they therefore incline to the hirsute covering of the muzzle of the Hip- popotamus.” In another placet the same author, speaking of the lifting muzzle of the upper lip (the levator labii superioris proprius), remarks, ‘“‘ Many vessels penetrate the root and origin of this levator ; * “ Tyansactions of the Zoological Society,” VIII, p. 133. t+ Lee. cit. p. 149. a ee Oe Beall —r — ON THE MANATEE. 305 this, no doubt, led Vrolik* to regard ‘the structure of the upper lip as plainly an erectile tissue.’ ’’ Observations on the living animal verify the correctness of Vrolik’s surmise. It is a post mortem change which causes the thickly bristled erectile lip-pads to be directed downwards. In the living animal their erectile fissue is distended with blood on most occasions, especially whilst feeding, and the pads are by this means directed inwards, towards one another, in such a way that the deep median notch which they go to form is even deeper and the bristles meet across the median line. Further, these pads have the power of transversely approaching towards and receding from one another simultaneously, Plate [11] XXVIII. figs. 1 and 2. When the animal is on the point of seizing, say a leaf of lettuce, the pads are diverged transversely in such a way as to Page 139. make the median gap of considerable breadth. Directly the leaf is within grasp the lip-pads are approximated, the leaf is firmly seized between their contiguous bristly surfaces, and then drawn inwards by a backward movement of the lower margin of the lip as a whole. The appearance produced by the movements of this peculiar organ is very much the same as that of the month in the silkworm and other caterpillars whilst devouring a leaf, the mandibles in these insect. larve diverging and converging laterally in a very similar manner during mastication. As to the mechanism of the process, the erectile nature of the lip- pads no doubt assists in the manner above indicated. These organs, when dilated, form more satisfactory muscularorigins than when flaccid. Innumerable transverse muscular fibres connect their basal portions, forming the bridge above the median lip-notch, and approximate them. They are evidently separated and diverged by the levator labii supe- rioris proprius, which is almost entirely distributed, in its insertion, to the considerable amount of skin-covered fibro-elastic tissue forming the interval between the nares and the lip-margin. . With reference to the valvular mechanism for closing the nostrils during submersion, it may be mentioned that these circular orifices have each a flap valve, which forms the floor or inferior wall of the nasal tubes when the animal is breathing, but which rises and com- pletely occludes it when closed, as represented in both the figures on Plate [11] XXVIII. Looking at the living animal generally, the most striking pecu- liarity was the sluggishness of its movements. When crossing its pond there was uone of the lateral movement of the body so charac- teristic of the Seals, All flexions were up and down, the whole trunk * “ Mem. Zoolog. Soc. Amsterdam,” 1852, p. 59. : x Page 140. 306 ON THE MANATEE. bending a little in that direction, the base of the tail doing so freely at a clearly marked transverse fold-line in that region. An opportunity occurred to me for seeing it out of water, when its pond was drained dry for a short time. From my observation on this occasion it is perfectly evident to me that the Manatee is purely aquatic in habits, and that it never willingly quits the water. When on land it seemed perfectly unable to advance or recede, the only movements it performed being that from its belly to its back, and wice versd. In these it made use of its limbs, flexing the body and the tail at the same time. When resting on its belly it seemed extremely uncom- fortable, apparently on account of difficulty in breathing, which it is easy to account for on the assumption that the weight of the body being supported on the almost ribless anterior walls of the thorax and abdomen, compressed the abdominal viscera against the lungs, and so greatly diminished the respiratory movements. It would not remain in this position for any length of time, but seemed comparatively comfortable whilst lying quietly on its back, as it did until the inflow- ing water had refilled the pond sufficiently to allow of its again supporting itself in the water. I may mention that the power of moving the slightly exserted elbow was considerable, whilst that of the wrist was small but appa- rent. It used its limbs much more freely than do the Seals, some- times employing the extreme margins of the paddles to assist in introducing food into the mouth, at others employing them in pro- gression along the bottom of the pond, during which time the swimming-tail could not be brought into play to any extent. The Manatee came into my hands within an hour or so of its death, and one of the earliest things that seemed desirable to do was to obtain some of its blood for examination. The first cut through the skin was sufficient to prove how different an animal it is from any of the Pinniped Carnivora; for instead of the muscles being of a deep almost black-red hue, as they are in the Seals, they were of a pale pink, more like veal or pork than any other flesh known to me, much lighter than beef. The blood-disks are circular and non-nucleated, as it was certainly known they would be. Their size, however, is their peculiarity. From the valuable investigations of Mr. Gulliver, which are incor- porated in their entirety in the Society’s ‘‘ Proceedings” for 1875 (p. 474 et seq.), it is known that the largest mammalian blood-disks are found in the Elephants (;,;; of an inch in diameter), Great Anteater (g747), Sloths (4/55), Aard-vark (5,4,,;), and Walrus (375). In the Manatee the diameter of the largest reaches 3755 of an inch, others being considerably smaller. If there is any stress to be laid on the size of the blood-disks in ON THE MANATEE. ; 307 the classification of animals, as it seems almost impossible that there should not be from the comparative constancy in their size in closely allied species and genera, then the relationships of the Manatee to the Artiodactylate Ungulates must be most distant, as small size of blood- disks is a special peculiarity of those latter animals, the largest being