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SENT Aan gate i 2: ‘aber arn 1 ae) Retaacy re AN oe aM WP Tea y ato ah Ue ath iin He th a i i oi im \e per rbhantih oe , yey vi Petit AK US eR AP ea Ae HR CITES CAN Hea Kan rans i A ty ty At La) a¥ anes 19s d's) AN eyeaoay eee iN (ahs) Vote bale VRC eva sole eee CAROR IR Mt OR MOUR EMI ION Wie} * iV winery pier yee COMME RE We 4 Hilateh Wah aig bicid AN A Pas t 3 x A vpn if TSS A HT NANT UA Or era un taid 8 i i oA, 74 « + ah Key! iy iar atb opt itive wean te vant ead et Mae tA Ma ge u Nt RYN A 4 Aid bet ia eS 8) iy We oimL AN NRA AL. | Dee ae ’ ORS ails Ds ie ue La WTNH a eRe) \ aN ASS aS Ny bee , Ay : 4] TE Mag hh vet Chatigny biel vex hat Naan ‘ en § ce) wk bs Wah Daath what! ; MCAS ae ote ee AE 4 * SOP a) ch ; Ss teas, Ave wee ory ve hi Pee ia ose Uc} ‘ Ue ey Weta a 9 TOC hy ‘ iuWe ne NE! 4 AA OCI Pav eval ut) aha afi’ east iy Ua a eK Oona) shy shy Pane \ ro Se) ‘ tag's 4 Verh ih) Re hence f) ee waa bos Pein ite SR REECE REA] Aas ‘ ae ZAR AWN eee ey yyy were) tayee COU ee) UNOS s PE Ne hire) fo) Wye Aas, ) a here 1k AR Vinee) ue vat SORA. ePN aan ke Vad ae Wy thick) i AM iNiey) Dat DUR EAN Ait Ri Seteentciie ty A OPTRA TAWA) he oe) bs oy OOOO POOR OLR IG MALY yyy Mite Witka AICI) aa { ‘a Wate LAUR LAD es (NW leral tea RASS Seeley me ahd OO wes ue Rich i ate iat uae i = ae at (<> [ 8B CHAPTER XI. GENERAL CONSIDERATIONS, : . : : . . > a los BIBLIOGRAPHY : - 5 . . 5 : - : oes) DeEscRIPTION OF PLATES . . . : 0 . : i ditesil INDEX : : 3 5 5 5 5 - ; a GR Snake loop Pps é Venom dried Soe Muscle tissue altered by venom 4. Muscle tissue altered by venom NR ORD UC ON: A rew words of explanatory character in regard to the following essay may not be out of place. From the time of Fontana, 1767, until the able essay of Lucien Bonaparte, in 1843, on the chemistry of venom, there was no paper of moment on serpent poisons. In January, 1861, one of us, S. Weir Mitchell, published a long study of the venom of the Crotalus durissus, and in 1868 sup- plemented it by a shorter contribution, in which he related some recent discoveries of his own, and corrected certain errors of his former paper. ‘These two essays may be considered as constituting with Lucien Bonaparte’s the foundation of the later work in this direction, and perhaps as having left the study of venoms in as definite a position as could be gained with the laboratory facilities of 1843 to 1868. In 1872, the government of India enabled Sir Joseph Fayrer to publish a volume of beautiful plates of the venomous snakes of India, to which was appended also a series of investigations into the toxicology of their poisons. In 1872 the same author and Dr. Lauder Brunton contributed an admirable physiological study of the effects of venoms.’ In 1874, Vincent Richards, as chairman of a government commission, published an excellent report on antidotes. Dr. Wall’s? thoughtful and suggestive book appeared in 1883. It is a compara- tive study of the poisons of the colubrine and viperine serpents of India. These, with a too brief study of the poison of our copperhead by Dr. Isaac Ott, of Easton, Pennsylvania, sum up all of value which has been added to the physiological. literature of this most interesting subject. Why it has won so few investigators is not far to seek. Even in India, where the appalling loss of life from snake-bites has of late invigorated research, the power and means of government were needed to overcome the obstacles which surround such scientific effort from inception to close. But, if in a land where snakes abound and professional snake-catchers can be had, it is yet not easy to follow this pursuit with success, elsewhere it is a task set about with inconceivable obstacles. The fear of serpents, the rarity of some species, the distances to which they have to be carried, the mortality of caged specimens, and the great cost of * Proc. Roy. Soc. 1872, 1873, and 1875. * Indian Snake Poisons; their Nature and Effects. A. J. Wall, M.D., F.R.Coll.S., 1883. all April, 1886. ( 1 ) wy) INTRODUCTION. purchase and transportation, need only to be mentioned as indicating our own difficulties. What had been done in India, sustained by a government, had to be with us attempted by private individuals, aided by the Smithsonian Institu- tion, without which it would have been impossible to succeed. Our work began in the autumn of 1882, by extended efforts on our part, and that of the Smith- sonian, to buy or otherwise get numerous living specimens of the American genera of Thanatophidee. This quest was kept up by every means our ingenuity could devise, and neither time nor money was spared. We succeeded in obtaining a sufficient number of rattlesnakes, including Crotalus adamanteus and C. durissus. We have had also enough of the Moccasin (Ancistrodon piscivorus). Our wants as regards Ground Rattlesnakes, Copperheads, and Coral-snakes have been less com- petently supplied, chiefly because these snakes are all small, so that to get enough of their poison for study it was essential to have a great many snakes. We have had in all about two hundred living serpents, and among them some superb specimens, which yielded poison in large quantities. Thus one—C. adamantews— was eight and a half feet long and weighed nearly nineteen pounds. It furnished on one occasion about one and a half drachms of venom. It was thought desirable by Prof. Baird and ourselves to examine the poisons of Indian serpents. ‘To secure these the Secretary of State appealed to Her Majesty’s Indian government in our behalf. A courteous response was returned, and orders given which resulted in our receiving a certain amount of Cobra venom. A more constant and larger supply was due to the generous and untiring kindness of Vincent Richards, Esq., M.R.C.S., of Goalundo, B. I. The poison of the Daboia Russellii, the Indian viper, we sought in vain to secure. Government aid and private enterprise alike failed to secure a sufficient quantity of the venom of this dreaded reptile. The other Thanatophidee, of Australia, and South America, still await more careful study, and our preliminary report has already been the means of renewing interest in the chemical aspects of this study in India. Such of our serpents as were not cared for by the hospitality of the Philadelphia Zoological Garden, were kept in large boxes, about four and a half feet high, covered on top with removable wire network, and well-ventilated through wired openings below. ‘They were of course furnished with water, and if they declined to eat, were fed at intervals, by artificial means, with raw beef chopped fine, and passed down into the belly of the snake through a large glass-tube. Under this treatment the deaths were fewer, and the supply of venom far better. Probably this method could be usefully employed in zoological gardens, where many snakes are lost owing to their indisposition to feed during the early months of captivity. On all occasions, for forced feeding, or for the purpose of extracting venom, the snakes were caught and held in the snake loop, Fig. 1. This is merely a staff, having a leather strap so arranged that it can be drawn out into a loop in which the serpent’s neck is noosed, and so held. With this simple means all risk is avoided, and with it serpents of any size and strength to be met with among our Thanatophidee can be safely held and easily manipulated. For whatever reasons the study of snake venoms had not greatly advanced since LENC eR ORD Ewe Cele ORN 33 the last research of Fayrer and Lauder Brunton until the authors of this paper resumed the work in 1882. One of them (Dr. Mitchell) had long felt that it would be well to revise the toxicology of our American serpents which he had begun in 1858, and as the later English observers had in some points differed from him, to learn if they or he were correct, or whether the divergence as to results was due to variations in the qualities of the venoms employed. Then too he had become conscious of certain errors in his former researches, and wished to aid in correcting them, and in filling up some of the gaps left in this branch of toxi- cology by himself and others. The authors started with a theory long held by Dr. Mitchell that snake venoms are not simple in composition, but composed of two or more poisonous substances, and that in the qualities and quantities of these agents would be found an expla- nation of the differences between serpent venoms as to power to kill and mode of causing death. How fertile has been the germinal idea of this research must be judged of by this present essay; which will, we trust, by leading thought and experiment in new directions hasten the day when we shall be able to treat with success the wretched thousands who now perish annually by snake-bite in India and elsewhere. Some of our earlier results were so soon talked of and even noted in public prints, that it seemed wise for this, and all other reasons, to state what we then knew. This was done in a “Preliminary Report to the United States National Academy of Sciences, in April, 1883.” In this brief essay we announced our proofs of the complex nature of snake poisons. ‘The report was incomplete, and in the light of our present more elaborate essay may be seen to contain several erroneous statements. It is not in the nature of things, that a research along such varied lines as our present volume follows, though extending over several years, should be per- fect in detail, or complete for all genera of Thanatophidians. It is our earnest 4 INTRODUCTION. hope that it will be complemented and supplemented by some of the able staff of the British Army Medical Service in the East Indies. ‘There, only, is it possible to find enough serpents, and all the various species which it will be desirable te review toxicologically from the new stand-point which we think we have estab- lished. We have forborne to overload this paper with comments on the later researches of others, and have made the discussion of our own work as brief as was consistent with clearness. In writing of the various substances contained in venoms, we have given them names which are fairly descriptive, but which, as in the case of the peculiar peptone of Cobra, may perhaps excite criticism. Yet, however unsatisfactory our method of nomenclature may be, any other plan of naming the curious bodies in question would certainly have been even more misleading. CHAPTER LI. PHYSICAL CHARACTERISTICS OF VENOM. Physical Characteristics of Venom.—All serpent venoms are more or less alike in appearance when fresh. ‘They are fluids varying in color from the palest amber tint to a deep yellow. Dr. Wall describes the Cobra venom as being occasionally colorless. ‘This peculiarity we have never seen in the fresh poison of any of our serpents, except once in the coral snake; nor can the venom of one kind of snake be distinguished with certainty by any physical peculiarity from that of any other, however remote they may be in the scale of being. When a fluid venom is allowed to dry slowly it presents no specific distinctive appearances. If desiccated too rapidly, it may look a little more gray and opaque than is common, but usually it dries into a beautifully cracked mass, deceptively like an aggregation of crystals, and which is well represented in Fig. 2. Fig. 2. In this state it is in solid yellow particles, very fragile, bright yellow, trans- parent or translucent, and seemingly indestructible by time, since the dried venom of the rattlesnake, for twenty-two years in Dr. Mitchell’s possession, proved as poisonous as that removed yesterday. It is equally unaltered by solution in glycerin, which keeps it permanently in unchanged toxic force, as we shall here- (5) 6 THE VENOMS OF CERTAIN THANATOPHIDE SA. after point out.! Neither does it appear to be injured when dry by mingling it with pure alcohol. In fact any of these three means, desiccation, glycerin, or alcohol, preserves it well. When fresh venom of any serpent is examined with the microscope it often presents a variety of floating bodies which seem to be much alike in all cases, and are very well shown in the plates of Dr. Mitchell’s former paper and in Vincent Richards’s reports. In healthy serpents, but lately caged, there are fewest of these solid ingredients, as has been noticed by Richards, by Wall, and by S. Weir Mitchell. ‘The question of the toxicity of these suspended solids has again drawn our attention to them, and we have had yet more careful and repeated microscopic examinations made by Prof. Formad. He found, like other observers, that the venom of the more vigorous snakes has the least visible solid matter; but, as in the use of the fang, the mucus and floating solids of the mouth must be considered, and, as in collecting venom from the snake, more or less of the mouth fluids mingle with the venom, it was thought well to reconsider the nature of the floating solids from the point of view of toxic activity. For the better study of the solids found in venoms we examined numerous specimens, and placed many of these in the hands of Prof. Formad, from whose notes we select the following observations :— A drop of fresh venom, taken directly from the Crotalus adamanteus, was examined with a =1, Zeiss. homog. immersion lens; amplification 800 diameters. The most striking appearance which first meets the eye is a granular material scattered about in masses of various size and shapes, resembling those formed by bacteria. ‘There are also seen, in some cases, a few oval nucleated red blood-corpuscles, some leuco- cytes resembling salivary corpuscles, and others corresponding to ordinary white blood-corpuscles, the latter cells in an active state of amceboid motion. There were also observed several club-shaped epithelial cells covered with fine granular material. The granular matter first mentioned, and which seems to form the main solid constituent of the venom, consists of two elements: Larger granules of an animal or albuminous character, and a fine granular material of vegetable nature. The albuminoid material is made up of minute particles ovoid, or somewhat irregularly angular in shape, measuring about ;54,, of an inch im their longest diameters. These ovoid particles are grouped side by side, from two to twenty in each collec- tion, and are arranged so as to form single or double rows, or more often aggregated into irregularly shaped clusters, which vary in size from =4, to ;j5, of an inch; the smaller masses predominating. The particles just described are colorless, refracting, and in general give the impression of bacteria. They are, however, distinguished from the latter in that they do not multiply in cultures, or respond to the aniline dye test for bacteria. There are usually numerous bacteria in perfectly fresh venom. All the smaller particles and granular material are micrococci, measuring on an average z>45> of 40000 an inch in diameter, are perfectly round or somewhat ovoid, and occurring singly, 1 Dr. Mitchell possessed a glycerin solution which was toxic after twenty years. PHYSICAL CHARACTERISTICS OF VENOM. 7 in pairs, or in zoogloea masses. They are less refracting, and paler than the albuminoid particles described above, and respond promptly to the usual tests for bacteria, viz: They multiply rapidly and absorb well the aniline dyes, thus form- ing a marked contrast side by side with the animal granular material, which was readily discolored under the influence of acid. The epithelial cells seen in the venom are, as a rule, few in number, are squam- ous or club-shaped, and in size not exceeding that of the red blood-corpuscle of the serpent. Leucocytes are also few in number, and, as well as the epithelium, are mostly covered with micrococci. A few of the white blood-corpuscles do not appear to contain micrococci, and in fresh venom, especially upon the warming stage, exhibit a quite active ameeboid motion. ‘The venom of the moccasin presents the same appearances. If fresh venom stands but a short time exposed to the air the micrococci mul tiply with remarkable rapidity, forming large, pale, motionless clonds; but, in addition, multitudes of movable bacteria (the Bacteriwm termo and a bacillus— probably Bacillus subtilis) gradually make their appearance.’ Tke globulous masses, above described, may be collected by filtration, but as this is often a difficult or even an impossible process with a fluid as viscous as pure venom, and, as much is lost in the filter, another method was devised, and there- after frequently used by us as an assistance in venom analysis. A tube, about 5 millimetres wide and 200 to 400 m.m. long, has a bulb blown on it midway, or at the top, and is then closed above in the blowpipe flame, and strongly heated throughout. While hot, the lower end drawn to a point, is in like manner sealed. After being cooled the tip is broken within fresh venom, which is forced up into the tube by atmospheric pressure. The end of the tube is then once more adroitly sealed in the flame. Thus prepared the tube is suspended, so that the solids of all forms settle in a few days, while for this time, at least, the venom undergoes no such putrefactive change as is inevitable when it is exposed to the air at our ordinary spring or summer temperatures. The solids, thus collected below, are easily separable from the supernatant venom by breaking off the two ends of the tube and allowing the precipitate to escape, with a minimum amount of liquid, from which washing in water easily separates them. The physical appearances of the venoms of the moccasin or of the rattlesnake, thus secluded from the air in these partial vacuum tubes, undergo some curious changes of much interest. The yellow coloring matter disappears from below upwards, and at last is seen only at the top, where the venom is in contact with the small amount of air left in the tube. At first, this change was presumed to be simply the rising of a pigment of lesser gravity. But it was noticed that the layer of yellow was of no deeper tint in its lessened bulk than when diffused. ‘The fluid below it was left as * Fresh venom, putrefied from long standing, appears to lose at least a portion of its virulence. But this is a point which is open to further observation. 8 THE VENOMS OF CERTAIN THANATOPHIDESA. clear and tintless as water; but when re-exposed to the air once more became yellow throughout, within one or two hours. The yellow pigment of Cobra poison, when the dry poison was dissolved in water, does not rise in the tube or disappear, but remains unaltered. It is desirable to repeat these observations with fresh Cobra venom. The cause of the disappearance and reappearance of the coloring matter of venom we have not been able to explain to our satisfaction, and it is one of the questions left open for inquiry. The Specific Gravities of Venoms.—The specific gravity of the venoms of our own serpents is as follows :— Crotalus horridus . : : : 3 2 : 5) JOR Crotalus atrox : : : A : 4 5 5 LoOre Crotalus adamanteus : $ - : : > UO Ancistrodon piscivorus . : : : : é . 1.032 The specific gravity of Cobra venom is given by Wall at 1.058. As to that of the Indian viper we can find no statement. The losses of venom on drying were as follows :— C. adam. : 0 . : : 5 . 25.15 per cent. C. atrox . , 3 F : ; : : 25.16 Ss Ancis. piscivorus . i f ; 3 Piteetey GG THE CHEMISTRY OF VENOMS g CHELA TPA Ui Ie THE CHEMISTRY OF VENOMS. Tue presence of alkaloids in venom, and especially of the ptomaines, has been suspected, and these bodies have been repeatedly sought for in vain. Gautier is the only chemist we recall who asserts that he found a ptomaine in a venom (Cobra). He does not state his processes, and we have been utterly unable to substantiate his statements. Lest we should in some way have erred in the con- duct of this part of our labor, we asked Prof. Wolcott Gibbs to examine Crotalus venom with a view to detection of such a body. As regards this search he makes the following statement :— “ My investigation of rattlesnake venom had for its special object the comparison of the venom with the higher alkaloids. As the quantity of material at my com- mand was small, I was obliged to content myself with the application of the ordi- nary tests used for the detection of alkaloids, as, for example, phospho-tungstates and phospho-molybdates, iodide of mercury and potassium, etc. etc. In many cases precipitates were obtained, but these were in no case distinctly crystalline. They resembled, on the contrary, the precipitate formed by sodic phospho-tungstate in solutions of albuminates in acetic acid. It seems, therefore, very improbable that the venom contains an alkaloid in the sense in which that term is commonly employed by chemists. On the other hand, it may still be basic in character, even if it be classed with albuminoids, since these are known to combine with platinous cyanide and with salicylic and other acids, exhibiting the properties of weak bases as well as of weak acids.” Venoms are of acid reaction, but when neutralized we have not observed any precipitate in specimens of these poisons. When venom is taken from the Crotalus or Ancistrodon there is often observed in the clear poison some insoluble whitish, granular matter, which soon settles to the bottom. The Insoluble Precipitate.—Vhis insoluble matter, which we term the insoluble precipitate, can be collected for examination by allowing the venom to stand in hermetically sealed vertical tubes, as previously described. The precipitate soon settles to the bottom, the clear venom is then carefully drawn off, and the precipi- tate is repeatedly washed with distilled water and collected; the washing process is repeated until there is no trace of proteid reaction in the wash-water, or, in other words, until all of the soluble portion of the venom has been completely washed from the precipitate. When examined under the microscope this precipitate consists of irregular 2 April, 1886. 10 THE VENOMS OF CERTAIN THANATOPHIDES. masses of granular matter with epithelial cells and salivary corpuscles, and a few flat crystals resembling cholesterin: The precipitate gives no proteid reactions with the usual proteid color tests, is insoluble in neutral saline solutions, and in weak or strong acids or alkalies. Boil- ing seems to render the mixture clearer. When injected into pigeons this precipitate does not appear to possess any toxic properties. Lhe Globulins.—If, after the separation of the above insoluble precipitate, the venom be mixed with water and placed in a dialyser over running water it will be found that within a few hours a whitish precipitate will occur within the dialyser, and should dialysis be continued sufficiently long the precipitate will have become deposited.in abundance. If the precipitate thus formed be collected on a filter it will be found that all of the coagulable proteids have been thrown down, since the filtrate now yields no coagulum by brief boiling, although it gives a proteid reaction. The precipitate is now washed from the filter and subjected to repeated wash- ings and decantations with distilled water, until the wash-water gives no proteid reaction. This purified precipitate is found to give reactions peculiar to the globulins ; it is insoluble in distilled water, soluble in dilute neutral saline solu- tions, soluble in dilute acids and alkalies, becomes turbid at about 60° C., and is fully coagulated at a point a little above 70° C. The filtrate still contains some proteid in solution, since we find, by the usual color and chemical tests, a proteid reaction, although it is observed that no coagu- lation occurs by momentary boiling. The filtrate is not precipitated by strong or weak: _ mineral acids, by solutions of ferric chloride or cupric sulphate, it is precipitated but not coagulated by absolute alcohol, and if placed in a dialyser it will be found to be readily dialysable. 'These reactions it will be observed place the proteid which remains in solution in the filtrate among the peptones. But we shall revert to this hereafter. It will thus be clear that we have separated in venom representatives of two distinct classes of proteids, one of which is insoluble in distilled water and coagu- lated in solution by boiling, and another which is soluble in distilled water and non-coagulable by brief boiling; the former belonging to the globulins and the other to the peptones. : The substance, however, which we find belonging to the globulins is a complex body in its composition, since, by appropriate processes, it can be resolved into three distinct principles, each of which is a globulin, but each having some properties different from its fellows. In order to distinguish these principles we have named them water-venom-globulin, copper-venom-globulin, and dialysis-venom-globulin, the names indicating the principal feature of the processes by which they are isolated from each other. As there are some differences in the reactions of similar prin- ciples in different species of venoms, we shall at first speak only of the venom of the Crotalus adamanteus. Water-venom-globulin.—We have already stated that when a solution of the fresh or dried venom in distilled water is allowed to stand for some time, especially if the quantity of water be comparatively large, a whitish precipitate occurs which THE CHEMISTRY OF VENOMS. il settles to the bottom of the glass, leaving in the course of a few hours a per- fectly clear supernatant liquid. If sufficient water has been added at first, the addition of more distilled water to the supernatant liquid will not cause any further precipitate. The precipitate is now collected and repeatedly washed with distilled water and decanted until the wash-water yields no proteid reaction. The following gives the results of some of the many reactions upon the addition of the various reagents used *—* Decided reactions with the usual proteid tests. Boiling—causes coagulation. Sodic chloride (0 75 per cent.)—slightly soluble. NOR cds )—soluble, forming a turbid solution ; the solution is not precipi- tated by carbonic acid? nor by the addition of ether. —boiling the solution causes coagulation. —the solution is precipitated by saturation with sodie chloride. Carbonic acid'—soluble. Sodie carbonate—very soluble; solution not precipitated by carbonic acid. Hydrochloric acid (0 4 per cent.)—very soluble. Metaphosphoric acid—insoluble. Orthophosphoric acid—dissolves. Sodic metaphosphate—insoluble. Sodie orthophosphate—very soluble. Potassic sulphate—very soluble. Calcic chloride—very soluble. Acetic acid (5 per cent. )—very soluble. Acetic acid (glacial)—very soluble. Coagulation occurs at about 64-73° C. Since this body is precipitated by saturation with sodic chloride, and dissclves with difficulty in a 0.75 per cent. solution of sodic chloride, it seems more akin to myosin than other of the globulins. The Copper-venom-globulin.—After the separation of the water-venom-globulin the filtrate gives well-marked proteid reactions and decided coagulation by boiling. If now a few drops of cupric sulphate (10 per cent.) be cautiously added a second precipitate will occur, and which can be separated as in the previous instance. In adding the cupric sulphate great caution must be exercised lest too much be added with the result of a complete or partial re-solution of the precipitate. The precipitate is sometimes comparatively slight at first, increasing upon stand- ing, and complete within about twenty-four hours. ‘The clear filtrate should give no precipitate after the addition of a small amount of the copper solution and after standing twenty-four hours longer. 1 Tn all of these reactions with the globulins, unless otherwise apparent, about 1 ¢. c. of the suspended globulin in distilled water was placed in a small test-tube, and from one to two drops of standard laboratory solutions of reagents were allowed to run down the inside of the tube. We have made a large number of tests with various reagents, and from this number have selected only such as will serve us some purpose in distinguishing these different bodies. 2 Where carbonic acid is used in these tests we have reference to the super-saturated carbonic acid water (soda water) of commerce. 12 THE VENOMS OF CERTAIN THANATOPHIDESA. The precipitate thus obtained is washed as in the preparation of the water- venom-globulin, and when thus purified it does not give any color reaction with the ammonia or the ferrocyanide and acetic-acid tests for copper, and therefore cannot be regarded as a salt of this metal. The copper-venom-globulin gives the following reactions: — Decided reactions with the usual proteid tests. Sodic chloride (0 75 per cent.)—insoluble. (CLO )—insoluble. —the addition of erystals of sodie chloride seems to dissolve it slightly; this solution is cleared somewhat by boiling; the same effect by boiling the suspended mixture; the clearing is no doubt the result of the formation of coagula. Carbonic acid—insoluble. Sodic carbonate—very soluble, forming a beautiful clear solution; boiling has no effect; the solu- tion is precipitated by carbonic acid. Hydrochloric acid (0.4 per cent. )—exceedingly soluble. Metaphosphoric acid—insoluble ; boiling no effect. Orthophosphoric acid—very soluble, forming an absolutely clear solution; boiling has no decided effect. Sodic metuphosphate—insoluble ; boiling no effect. Sodic orthophosphate—soluble in a much larger amount than is necessary in dissolving the water- venom-globulin; boiling has no effect, unless to clear the solution some. Potassic sulphate—insoluble; boiling no effect. Calcic chloride—less soluble than water-venom-globulin. Acetic acid (5 per cent.)—very soluble. Acetic acid (glacial)—very soluble. The Dialysis-venom-globulin.—The filtrate, after the separation of the water- venom-globulin and copper-venom-globulin, still gives a decided amount of coagula by boiling, and also all of the characteristic color reactions for proteids. If the filtrate be now subjected to dialysis, best by means of a large dialyser placed over running water, in the course of twenty-four hours a considerable amount of pre- cipitate will be deposited within the dialyser, and which may be collected on a filter, and repeatedly washed as in the preparation of the preceding globulins. If dialysis is carried on for a sufficient length of time the whole of this principle will be precipitated, since the filtrate from the globulin will give no coagula by boiling, nor any precipitate by strong nitric acid. A proteid still remains in solu- tion, however, which has been already alluded to as being a peptone. ‘This body being less dialysable than the salts which hold the globulins in solution, still remains in part within the dialyser, even when the salts are so fully withdrawn as to entirely precipitate the globulins. The dialysis-venom-globulin gives the following reactions :— Decided reactions with the usual proteid tests. Sodie chloride (0.75 per cent. )—insoluble. GO. )—slightly soluble. (erystals)—more soluble, forming a very cloudy solution; boiling clears the solution some; the same degree of clearing does not occur in the mixture without the sodic chloride. —the addition of carbonic acid to the solution with erystals causes a beautiful clear solution, which is made cloudy by boiling. THE CHEMISTRY OF VENOMS. 13 Carbonic 2cid—soluble; cloudiness by boiling. Sodic carbonate—very soluble ; boiling no effect. Hydrochloric acid (0.4 per cent.)—very soluble. Metaphosphoric acid—rendered of a yellowish tint; not appreciably dissolved ; boiling no appre- ciable effect. Orthophosphoric acid—very soluble ; boiling no effect. Sodic metaphosphate—very soluble, forming a very clear solution ; boiling no effect. Sodic orthophosphate—slightly soluble; dissolving slowly in excess, forming a slightly turbid solution ; boiling clears absolutely. Potassic sulphate—insoluble ; boiling no decided effect. Calcic chloride—soluble by the addition of a comparatively larger amount; boiling causes coagulation. Acetic acid (5 per cent.)—very soluble. Acetic acid (glacial)—very soluble. The Venom Peptone.—After the separation of the dialysis-globulin the filtrate, as before stated, gives no coagula by brief boiling, but by testing with the usual proteid tests very decided reactions are obtained. It is further found that if the above filtrate is placed in a fresh dialyser, that the principle giving the proteid reactions will readily pass through the membrane. ‘The fact that this substance will dialyse readily, and that it is not immediately coagulated at the temperature of boiling water, and not precipitated by cupric sulphate and ferric chloride, nor by neutrali- zation, renders it certain that it belongs to a peculiar class of bodies which are known as peptones, and which are ordinarily the result of peptic or tryptic digestion, ‘This peptone may also be prepared by briefly boiling the solution of yenom, which coagulates the other albuminous principles, and leaves this in solu- tion; but the coagula caused by boiling the solution of Crotalus are so extremely fine, that it is impossible to filter the mixture clear, even by repeated filtration through many thicknesses (7) of the best filter paper ; furthermore, continued boil- ing causes a breaking down of the peptone with the apparent formation of fine coagula (see Cobra peptone, p. 17). We, however, prepared the peptone by dialysis, and obtained the following reactions :— No immediate coagulation at a temperature of 100° C. Full reactions with the proteid color tests. No precipitate with weak or strong nitric acid. Ferric chloride—no precipitate. Cupric sulphate—no precipitate. Mercuric chloride—decided precipitate. Absolute alcohol—precipitate ; precipitate redissolved by the addition of water. Mercuric nitrate—decided precipitate. Potassic hydrate—precipitate by saturation; precipitate redissolved by the addition of nitric acid, forming a decidedly yellowish solution, which becomes decolorized by further addition of acid. Potassie ferrocyanide in presence of weak acetic acid—a precipitate. To revert now to the globulins and their distinctive features, it seems clear that these principles must exist in the venom as distinct bodies, and are not simply representatives of a single globulin which have arisen through our manipulations. The first distinguishing feature between them is represented in the process of isolation, but if we place the reactions of the different globulins in parallel columns, 14 THE VENOMS OF CERTAIN THANATOPHIDE A. we find that, while they have very close resemblances, as they naturally should since they are so intimately related, they are very readily distinguished from each other. The properties of all globulins are so readily affected by even the simplest manipu- lations that it is likely that mere precipitation may affect them in regard to their solubility, while drying may completely destroy this property. Having these facts in mind, it seems almost a necessity that the processes through which we put these globulins, in order to get them isolated in a pure state, has more or less modified their chemical, and possibly their physiological properties. The tests made with these globulins were all made at different times, the one globulin was examined one day, and another on another day, so that the reactions given are not absolutely accurate as a matter of comparison, but only relative, since the standard of solubility, which was of course an arbitrary one, was simply carried in the mind throughout the examinations. practically correct. We believe, however, that they are Reagent. Water-yenom-globulin. Copper-venom-globulin. Dialysis-yenom-globulin. Sodic chloride (10 p. ec.) Carbonic acid Sodic carbonate Hydrochloric acid (0.4 p. ce.) Soluble Soluble (Very soluble; not (precipitated by CO, Very soluble Insoluble Insoluble Very soluble ; pre- )} cipitated by CO, 5 Very soluble Slightly soluble. Soluble. Very soluble. Very soluble. (Insoluble; rendered Metaphosphoric acid Insoluble Insoluble alia gellowan omar Orthophosphoric acid Soluble Very soluble Very soluble. Sodic metaphosphate Insoluble Insoluble Very soluble. Sodic orthophosphate Very soluble Less soluble Still less soluble. Potassic sulphate Very soluble Jnsoluble Insoluble. Calcic chloride Very soluble Less soluble Less soluble. Acelic acid (5 per cent.) Very soluble Soluble Very soluble. Acetic acid (glacial) Very soluble Soluble Very soluble. The venom of the Moccasin (Ancistrodon piscivorus) was subjected to an analysis similar to that of the Crotalus, the isolated proteids giving the following reactions :— Water-venom-globulin. Decided reactions with the usual proteid color tests. Boiling—clears the mixture without the apparent formation of any coagula. Sodic chloride (0.75 per cent. )—insoluble. (Oe as )—somewhat soluble, solution not absolutely clear; boiling clears absolutely without the formation of coagula. (crystals)—somewhat soluble; solution not precipitated by carbonic acid. Carbonic acid—insoluble. Sodie carbonate—soluble, forming slightly turbid solution; boiling clears the solution without giving coagula; the addition of erystals of sodie chloride to the hot boiled solution causes a precipitate, this precipitate being coagulated by boiling. Hydrochloric acid (0.4 per cent.) —somewhat soluble. (Gries )—soluble. Metaphosphoric acid—insoluble. Orthophosphoric acid—soluble. Sodie metaphosphate—slightly soluble; solution rendered clearer by boiling. Sodic orthophosphate—soluble; solution rendered absolutely clear by boiling. THE CHEMISTRY OF VENOMS. 155 Potassic sulphate—soluble ; solution rendered absolutely clear by boiling. Calcic chloride—soluble ; solution rendered clearer by boiling. Acetic acid (5 per cent.)—insoluble. Acetic acid (glacial)—insoluble ? Copper-venom-globulin. Boiling—clears somewhat; no coagula. Sodic chloride (0.75 per cent. )—insoluble. GO )—insoluble. (crystals)—insoluble; boiling partially clears without the formation of any coagula. Carbonic acid—somewhat soluble; boiling clears absolutely. Sodic carbonate—very. soluble; boiling no effect. Hydrochloric acid (0.4 per cent.)—very soluble. Metaphosphoric acid—insoluble; boiling appears to clear slightly. Orthophosphoric acid—very soluble. Sodic metaphosphate—insoluble; boiling clears somewhat. Sodic orthophosphate—somewhat soluble; boiling clears beautifully. Potassic sulphate—insoluble ; beiling clears slightly. Calcie chloride—slowly dissolved; not so soluble as water-globulin; boiling gives a slight cloudiness. Acetic acid (5 per cent.)—soluble. Acetic acid (glacial)—soluble. Dialysis-venom-globulin. Boiling—clears almost absolutely without the apparent formation of coagula; boiled solution precipitated by saturation with crystals of sodic chloride. Sodic chloride (0.75 per cent.)—insoluble. (@i0s )—somewhat soluble; dissolves slowly, forming a slightly turbid solution; boiling seems to clear some without the formation of coagula. Carbonic acid—very soluble; slight turbidity by boiling. Sodic carbonate—very soluble; boiling no effect. Hydrochloric acid (0.4 per cent.)—very soluble. Metaphosphoric acid—slightly soluble; yellowish tint; boiling clears slightly with the formation of coagula. Orthophosphoric acid—very soluble ; boiling no effect. Sodic metaphosphate—insoluble. Sodic orthophosphate—soluble ; boiling no effect. Potassic sulphate—somewhat soluble. Calcic chloride—very soluble, form a beautiful clear solution; boiling causes slight turbidity. Acetic acid (5 per cent.)—soluble Acetie acid (glacial)—soluble Moccasin Peptone. 1. Readily dialysable. 2. Not immediately coagulated at a temperature of 100° C., but gradually coagulated by pro- longed boiling (see Cobra peptone, p. 17). 3. Reaction with the xantho-proteic test (nitric acid and ammonia) 4. Reaction with Millon’s reagent (mercuric nitrate) 5. No precipitate with weak or strong nitric acid. 6. No precipitate with CO,. 7. No precipitate with ferric chloride. 8. No precipitate with cupric sulphate. Gi THE VENOMS OF CERTAIN THANATOPHIDEA. 9. Precipitated by mercuric chloride. 10. Precipitated by absolute alcohol. 11. Gives a faint reddish tinge with a strong solution of potassium hydrate, and a trace of cupric sulphate. 12. Not precipitated by strong acetic acid (glacial). 13. Precipitated by very dilute acetic acid; precipitate being redissolved by further addition of acid. 14 Full reaction with Adamkiewicz’s test for proteids. 15. Precipitated by adding a large quantity of sodium chloride, the precipitate being redissalved on the addition of a large quantity of glacial acetic acid. 16. Precipitated by mercuric nitrate. 17. Precipitated by absolute alcohol; precipitate being apparently redissolved on the addition of water. 18. Precipitated by saturation with potassium hydrate; precipitate being redissolved by the addi- tion of nitric acid, with the formation of a decidedly yellow solution (xantho-proteic) which becomes decolorized by addition of acid. 19. Precipitated by potassium ferrocyanide in the presence of weak acetic acid. Venom-peptone by dialysis vives identical reactions. to) For convenience of comparison we append here in parallel columns the principal reactions of the Moccasin globulins, remembering in this connection the difference in the properties manifest in their methods of preparation. Reagent. Water-venom-globulin. | Copper-venom-globulin. | Dialysis-venom-globulin. Boiling eee los: Clears some Clears some. Sodic chloride (10 per cent.) | Somewhat soluble Insoluble Somewhat soluble. Carbonic acid Insoluble Somewhat soluble Very soluble. Sodic carbonate Soluble Very soluble Very soluble. Hydrochloric acid (0.4 p.c.) | Somewhat soluble Very soluble Very soluble. Metaphosphoric acid Insoluble Insoluble Slightly soluble. Orthophosphoric acid Soluble Very soluble Very soluble. Sodic metaphosphate Somewhat soluble Insoluble Insoluble. Sodic orthophosphate Soluble Less soluble Soluble. Potassie sulphate Soluble Insoluble Slightly soluble Calcie chloride Soluble Insoluble Very soluble. Acetic acid (5 per cent.) Insoluble Soluble Soluble. Acetic acid (strong) Tnsoluble Soluble Soluble. For reactions of the peptones of the various venoms see p. 19. Cobra Venom.—We have been able to isolate in Cobra venom only two proteids, and these correspond in their characters to the two types of proteids found in the venoms of the Crotalus and Ancistrodon. In other words, we have isolated a globulin and a peptone-like principle. The globulin we are able to precipitate completely by the addition of a proper amount of distilled water, after which the solution gives no coagulum by boiling. ‘There is then left in solution a proteid, which evidently belongs to the peptones, although giving some extraordinary reactions. The venom-globulin thus isolated and purified, as in the preparation of the globulins previously mentioned, possesses the peculiar properties of the globulin family, and, in accordance with our nomenclature, since it is entirely precipitated by the addition of distilled water, is a water-venom-globulin. THE CHEMISTRY OF VENOMS. 17 The following are some of the reactions given by this substance (the water-venom- globulin suspended in distilled water) :— Boiling—coagulates. Sodic chloride (0.75 per cent. )—insoluble. CO )—soluble; boiling gives slight turbidity. —sodic chloride solution apparently unaffected by carbonic acid. Carbonic acid—insoluble. Sodic carbonate—soluble, slightly turbid solution; boiling makes perfectly clear. Hydrochloric acid (0.4 per cent.)—soluble. Metaphosphoric acid—insoluble; boiling no appreciable effect. Orthophosphoric acid—very soluble; boiling makes solution absolutely clear. Sodic metaphosphate—insoluble; boiling no appreciable effect. Sodic orthophosphate—somewhat soluble; boiling renders perfectly clear. Potassic sulphate—somewhat soluble. Calcie chloride—soluble; opalescence of solution increased by boiling. Acetic acid (5 per cent.)—soluble. Acetic acid (glacial)—soluble. Cobra-venom-peptone.—The venom-peptone from Cobra may be prepared by boiling, thus coagulating the globulin, or by dialysis. Great difficulty is expe- rienced in the former process, since the coagula are so fine that it is impossible, save in rare instances, to obtain a clear filtrate, and as to these we have no explana- tion to offer for the exception. ‘The peptone prepared by boiling or by dialysis gives identical reactions. Before detailing the reactions of this body it may be well to notice a peculiar property exhibited by all venom-peptones which gives them a very distinguishing feature. After boiling the venom for a few minutes and then filtering, the filtrate will again give further coagula by continued boiling, and so the process of boiling and filtering, and reboiling the filtrate may go on repeatedly, yet the clear filtrate will in every instance give fresh coagula. Indeed the boiling process may be con- tinued for an hour or more, and yet at the end of that time the filtrate will still yield coagula. However, after the venom solution has once been boiled, coagula- tion does not recommence in the filtrate until it has been boiled for a few moments. These most interesting facts suggest that the coagula formed after the first boiling are due to a gradual decomposition of what is in some sense a non-coagulable pro- teid, since coagulable proteids all coagulate at once and completely when a definite temperature is reached; the coagula which follow repeated or prolonged boiling appear to be due to such a decomposition of proteids as violent chemical or physi- cal action could alone account for. It seems to us perfectly clear that the body which is thus gradually broken up by prolonged boiling is a peptone. Our principal reasons for this belief are that the body so coagulated is very readily dialysable, is not precipitated by ferric chloride, or cupric sulphate, and in the case of the Cobra is not precipitated by abso- lute alcohol, or mercuric chloride, is not coagulated below the boiling point, and in fact not until boiling has gone on for a few moments. ‘The following reactions seem to be sufficiently characteristic. These results we obtained from a solution of the Cobra-venom-peptone obtained 3 April, 1886. 18 THE VENOMS OF CERTAIN THANATOPHIDE SA. by dialysing venom for forty-eight hours. The dialysate was perfectly clear and neutral in reaction :— Boiling—no result until after a few moments, when it becomes cloudy, the cloudiness increasing as boiling continues; strong nitric acid dissolves the precipitate. Color reactions for proteids—the xantho-proteic, Millon and Biuret reactions are all obtained. Ferric chloride—no effect. Cupric sulphate—no effect. Mercurie chloride—no effect. Mercurie nitrate—decided precipitate. Absolute alcohol—no precipitate. Potassice ferrocyanide + weak acetic acid—precipitate. Nitric acid (strong)—no precipitate. Hydrochloric acid (strong)—no precipitate. Acetic acid (strong)—no precipitate. Sodic chloride (saturation)—precipitate; acetic acid, large quantity, dissolves. Potassic hydrate to satwration—precipitate. Tannie acid—decided precipitate. Basie acetate of lead—decided precipitate. Several very remarkable facts are the coagulation by prolonged boiling and the non-precipitation by mercuric chloride and absolute alcohol. Since this peptone is precipitated by weak acetic acid in the presence of potassic ferrocyanide it has a slight resemblance to Meissner’s A peptone, although materially differing, as some of the above reactions show, from any other described body of this class. As a matter of some interest, it is desirable to know if similar globulins in different venoms are identical in their chemical nature, or whether they give any reactions which may distinguish them. We have accordingly, as in previous cases, placed the reactions of the corresponding globulins side by side. I. Water-venom-globulin. Reagent. Crotalus horridus. Ancistrodon piscivoris. Cobra. Boiling Coagulates Apparently dissolves Coagulates. Sodic chloride (10 per cent.) Soluble Somewhat soluble Soluble. Carbonic acid Soluble Insoluble Insoluble. Sodic carbonate Soluble Soluble - Soluble. Hydrochloric acid (0.4 p. ¢.) Soluble Somewhat soluble Soluble. Metaphosphoric acid Insoluble Insoluble Insoluble. Orthophosphoric acid Soluble Soluble Soluble. Sodic metaphosphate Insoluble Somewhat soluble Insoluble. Sodic orthophosphate Very soluble Soluble Somewhat soluble. Potassie sulphate Very soluble Soluble Somewhat soluble. Calcic chloride Very soluble Soluble Soluble. Acetic acid (5 per cent.) Soluble Insoluble Soluble. Acetic acid (strong) Soluble Insoluble Soluble. THE CHEMISTRY OF VENOMS. 19 Il. Copper-venom-globulin. Reagent. Crotalus horridus. Ancistrodon piscivorus. Boiling Sodic chloride (10 per cent.) Carbonic acid Sodic carbonate Hydrochloric acid (0.4 p. ¢.) Metaphosphoric acid Orthophosphoric acid Sodic melaphosphate Sodie orthophosphate Potassie sulphate Calcie chloride Acetic acid (5 per cent.) Acetic acid (glacial) Coagulates Insoluble Insoluble Very soluble Very soluble Insoluble Very soluble Insoluble Soluble Insoluble Soluble Soluble Soluble Apparently dissolves. Insoluble. Somewhat soluble. Very soluble. Very soluble. Insoluble. Very soluble. Insoluble. Soluble. Insoluble. Insoluble. Soluble. Soluble. Ill. Dialysis-venom-qlobulin. ~ Reagent. Crotalus adamantcus. Ancistrodon piscivorus. Boiling Coagulation No coagulation ? Sodic chloride (10 per cent.) Somewhat soluble Somewhat soluble. Carbonic acid Soluble Very soluble. Very soluble. Very soluble. Slightly soluble. Very soluble. Sodie carbonate Very soluble Hydrochloric acid (0.4 p. ©.) Very soluble Metaphosphoric acid Insoluble Orthophosphoric acid Very soluble Sodic metaphosphate Very soluble Insoluble. Sodic orthophosphate Soluble Soluble. Potassic sulphate Insoluble Slightly soluble. Calcie chloride Soluble Very soluble. Acetic acid (5 per cent.) Soluble Soluble. Acetic acid (glacial) Soluble Soluble. It will be noticed by a careful comparison that the corresponding principles in different venoms differ quite as much from each other as the globulins in any one variety of venom. Venom Peptones.—We have not been able to detect any chemical differences in the venom peptones of the Crotalus and Ancistrodon. Cobra venom peptone is distinguished from that of the Crotalus and Ancistrodon by its non-precipita- bility by mercuric chloride and absolute alcohol. Daboia Venom.—We have had a small quantity (a few grains) of Daboia venom at our disposal, but too little to attempt any detailed chemical investigations. In two examinations, however, with very small quantities, we separated two bodies corresponding to those in Cobra, that is a water-venom-globulin and a peptone. The former exists in exceedingly small quantity while the latter dialyses with appa- rently much more difficulty than that of the Cobra. The Proportions of Proteid Constituents in Different Venoms. An examination of good specimens of the dried venoms of the Crotalus adamanteus, Ancistrodon piscivorus, and Cobra gives us the following proportions of the globulins and peptones :—— 90, THE VENOMS OF CERTAIN THANATOPHIDES. Crotalus adamanteus— 0.5 gram dried venom = water-venom-globulin 0.0495 copper-venom-globulin 0.0375 dialysis-venom-globulin 0.0360 0.1230 = globulins. 0.3770 = peptone’ (estimated. ) Ancistrodon piscivorus— 0.3364 gram dried venom = water-venom-globulin 0.0034 copper-venom-globulin 0.0182 dialysis-venom-globulin 0.0047 0.0263 = globulins. 0.3101 = peptone’ (estimated). According to this estimate there would be in 0.5 gram 0.0391 globulins. 0.4609 peptone." Cobra— 0.2 gram dried venom = water-venom-globulin 0.0035 peptone’ 0.1965 (estimated). According to this estimate there would be in 0.5 gram 0.0087 globulin. 0 4912 peptone.’ From these analyses it will be observed that the dried venom of the Crotalus adamanteus contains 24.6 per cent. of globulins, the Ancistrodon 7.8 per cent., and the Cobra 1.75 per cent. The globulins in the Crotalus venom appear to be in almost equal proportions, while in the Ancistrodon the copper-venom-globulin is about five times greater than the water-venom-globulin and about four times more than the dialysis-venom-globulin—the two latter being nearly in the same propor- tion—therefore constituting more than half of the entire weight of globulins, These differences in the proportions of the various globulins in any specimen of venom and the differences in the proportions of globulins and peptones in different venoms are of immense importance in affording an explanation of the physiological peculiarities exhibited in poisoning by different species of snakes. It will be observed that the proportion of globulins in Crotalus is over three times the quantity in the Ancistrodon, and nearly fifteen times that in the Cobra. 1 Including the salts, which are in very small quantity. EFFECTS OF VARIOUS AGENTS ON VENOM. 91 CHHVAGE Tek Resell EFFECTS OF VARIOUS AGENTS ON VENOM.. Liffects of Various Agents on Venom.—The influence of acids, alkalies, and salts on venoms has been studied by several observers, with results which vary remark- ably; so that for this and for other reasons there is still room for research of this nature. The questions thus brought up have a twofold interest, the one chemical and the other toxic. Numerous bodies precipitate or dissolve venoms; but among those which most plainly alter these poisons, only a few so change them as to lessen or destroy their poisonous efficiency. Unfortunately, that which alters the poison as such, is always equally destructive to the tissues of the body, and no agent as yet employed can be shown to have the power to enter the blood, and there affect the venom without doing harm to other albuminous substances. So far, we have learned only that amidst the agents which precipitate venom, there are some which weaken or annihilate its toxic force. They can be thrown into the fang tracks, and where they are made to mingle with the venom will destroy it as impartially as they do the mnocent tissues in which it les. It may not be out of place to remark that we have made no direct study of agents as antidotes. ‘loo much yet remains to be known of these poisons before we can hope to find a means of antagonizing them physiologically. Our local or chemical antidotes are sufficiently effective. Effect of Desiccation of Venom.—Allowed to dry at ordinary temperatures, the venoms retain their poisonous activity almost unaltered. When again water is added they act as usual, except that, owing perhaps to imperfections in redissolu- tion, they do not produce as much local effect within as short a time as do the fresh fluid venoms. Neither, it may be added, is the general toxic influence quite as rapid when venom has been once desiccated. The Effects of Various Agents on the Toxicity of Venoms. Age.—Some fresh venom of the Crotalus horridus was dissolved in an equal quantity of pure glycerine and the vial corked and sealed in 1863. In November, 1882, the contents of the vial were examined. The solution was perfectly clear, and had at the bottom a small mass of what appeared to be a fungous growth. Some of the venom was now injected into various animals to test its toxicity. The following experiment attests its power :— Experiment.—Pigeon. Injected, at 5:12 P. M., into the muscles of the thigh about six drops of the above glycerin solution. 5:14, Animal decidedly weakened. 5:25. There is considerable blackening of the tissues about the point of injection, the parts 22 THE VENOMS OF CERTAIN THANATOPHIDESA. being much swollen, the leg stiff, the muscles at the point of injection are paralyzed, and sensibility of the leg destroyed. ‘The pigeon lies on its side unable to stand, is exceedingly prostrated, and breathes laboriously. Observation now ceased until 8 A. M. following morn- ing, when the animal was found dead and in general rigor mortis, excepting the muscles at point of injection. Autopsy.—The tissues were dark, congested, and suffused with serum for an area of one and a half inches from point of injection. he viscera of the thoracic and abdominal cavities appeared slightly congested; the heart was arrested in systole and contained dark clots; the blood everywhere was dark and clotted. Microscopically the muscular fibres did not appear to be greatly disorganized, although in some of the fibres no transverse strie or nuclei could be discovered. The Effects of Dry Heat. Experiment.—0.03 gram of dried (Crotalus adam- anteus) venom was subjected in a dry oven to a gradually rising temperature to 83.5° C., and maintained at this point for half an hour. The venom, after cooling, was dissolved in 1 ¢.c¢. of distilled water. 2:57. Injected the above into the thigh of a pigeon. 4:49. Violent convulsions and death. Local effects decidedly marked. Experiment.—Repeated the above, but subjecting the venom to a temperature of 100 C. for ten minutes. 3:42. Injected into the thigh of a pigeon. 6:00. No decided symptoms. On the following morning the animal was dead. The local effects were marked. Experiment.—Repeated the above, but subjecting the venom to a temperature of 110° C. for thirty minutes. 4:46. Injected into the thigh of a pigeon. 5:25. Convulsions. 5:45. Died. The local effects were marked. From these results it seems clear that heating the dry venom to a degree above boiling point daes not apparently alter its poisonous activity. The delay in the occurrence of death in the second experiment suggests that the venom was altered, but in the third experiment in which the temperature was even higher, and this degree of heat maintained for a much longer time, death occurred even sooner than in the first experiment, showing that the differences must have been dependent upon conditions in the animals. The Effects of Moist Heat. Experiment.—0.03 gram dried venom (Crotalus adamanteus) was dissolved in 1 c.¢. distilled water, and gradually heated until a flocculent precipitate occurred. This was injected into the thigh of a pigeon in the evening. The next morning the animal was found dead. : Experiment.—0.03 gram dried venom (Crotalus adamanteus) was dissolved in 1 c. c. distilled water and subjected to a gradually rising temperature to 50° C. 3:49. Injected the above into the breast muscles of a pigeon. 3:51. Very weak, pupils apparently contracted, trembling; breathing laborious. 4:00. Dead. At the point of injection the tissues were decidedly congested and purplish and suffused with blood. The blood generally was fluid, but some soft clots were found in the abdominal vessels. EFFECTS OF VARIOUS AGENTS ON VENOM. 93 Experiment.—Subjected a similar amount of venom in solution to a rising temperature to 65° C. 4:01. Injected into the breast muscles of a pigeon. 4:05. Head depressed. 4:10. Very weak, falls on the side. 5:02. Dead. The local effect is not so marked as in the previous experiment. The injec- tion was merely subcutaneous. The viscera did not appear congested or abnormal; the heart was arrested in systole; blood everywhere fluid and dark; no ecchymoses in the peritoneum ; muscles appear darker than normal. Experiment.—Repeated the above, but increasing temperature to 74° C. 4:13. Injected into the breast muscles of a pigeon. 4:19. Weak, falls on side. 5:05. Dead. Blood clotted; local effect the same as in previous experiment. Experiment.—Results the same as in the last experiment, excepting that the local effects were more marked. ‘This animal lived a half hour longer than the last, which will probably account for the difference. Experiment.—The same, but subjecting the solution to 76.5° C, 4:04. Injected into the breast muscles of a pigeon. 4:27. Unable to stand. 6:00. Nearly dead. Following morning. Extremely feeble, too weak to stand; there is a muco-sanguinolent discharge from the bowels. Second day. Very feeble. Third day. Recovering. Experiment.—The same, but subjecting the solution to 79.5° C. 4:00. Injected into the breast muscles of a pigeon. 5:50. No symptoms. Following morning animal well. Experiment.—The same, but subjecting the solution to 81° C. 4:31. Injected into the breast muscles of a pigeon. 4:45. Apparently a little stupid. 5:50. No further effect. Following morning. Animal well. Second morning. Animal well. Experiment.—Boiled a similar amount of solution for two minutes. 3:26. Injected into the breast muscles of a pigeon. 4:30. No effect. Following morning. No effect. The above very interesting series of experiments clearly shows that the effect of heat on a solution of venom is very positive, that the toxicity of venom is decidedly affected, and that the greater the increase of temperature between certain limits the greater is the destruction of the poisonous power of the venom. It will be observed in the second experiment, which is the first in which any positive temperature was observed, that the animal died in about ten minutes after injection; in the third experiment in about one hour ; in the fourth and fifth experiments in about three-fourths of an howr, and an hour and three-quarters respectively; in the sixth experiment in about two hours ; the animal was nearly 94 THE VENOMS OF CERTAIN THANATOPHIDE A. dead, but finally recovered in the seventh experiment and in the subsequent ones there were no poisonous symptoms. It will thus be observed that there is a gradual impairment of the toxicity of the venom increasing with the increase of tempera- ture, and that when we reach 76.5° C. we have almost reached the temperature at which toxicity seems to be completely destroyed. We say seems completely destroyed, because we have found that the solution is still toxic even when boiled, although there is not sufficient active poisonous matter left after boiling in the small amount of venom we used in this group of observations to cause decidedly poisonous effects im pigeons. The results of boiling solutions of Moccasin and Cobra venoms are quite different from the above, as the following experiments clearly show:— Experiment.—Dissolved 0.015 gram dried Moceasin in 1 c. e. distilled water, and gradually heated to 78° C. 3:40. Injected into the breast of a pigeon. 3:50. Rocking. 4:45. Nearly gone; some local effect. Following morning the animal was dead. The local effect (darkening) was marked, but not comparable to that caused by the unboiled venom. Experiment.—Boiled 0.015 gram dried Moccasin (piscivoris) dissolved in 1 ¢. ec. distilled water for one minute. 3:28. Injected the above into the breast muscles of a pigeon. 3:35. Too weak to stand. 4:15. Dead. ‘There are no local effects. Experiment.—Dissolved about 15 minims of fresh Moccasin venom in about 1 c. c. distilled water, then boiled in a test-tube, filtered and injected one-half into the breast muscles of a pigeon at 4:30. 4:55. Very slight local effect; darkening and swelling; the animal is weak and has respi- ratory disturbance. Injected the other half. 5:00. Rocking; irregular breathing ; somewhat stupefied. 5:20. Hyes closed; stupefied; breathing irregular. Following morning. There was a large, light-colored, edematous swelling (see Plate No. 1) within, which was a cavity about an inch in diameter, full of broken-down tissue, having a grayish muddy, gangrenous appearance, and a putrefactive odor, while the surrounding muscular tissues were normal in appearance. It will be observed in this series of experiments with the Moccasin venom that there is also a very decided alteration in the poisonous properties of the venom. But here we find that although the amount of venom used was only one-half the quantity employed in the Crotalus series, boiling does not destroy its ability to kill. It will also be noticed here, as in the case of the Crotalus, that a sufficient degree of heat has an obvious effect on the power of the venom to produce the peculiar lesions at the point of injection. The effect of heat upon solutions of Cobra venom is not so marked. Experiment.—0.03 gram dried Cobra venom was dissolved in 1 c.c. distilled water and subjected to a temperature gradually rising to 74° C. 4:10. Injected into the breast muscles of a pigeon. 4:16. Unable to stand. 4:20. Dead. EFFECTS OF VARIOUS AGENTS ON VENOM. 25 Eaperiment.—The same, excepting that the temperature was raised to 79.5° C. 4:12. Injected as above. 4:21, Unable to stand. 4:25. Dead. Experiment.—The same, solution being brought to boiling point in a test-tube. 4:45. Injected into the breast muscles of a pigeon. 5:00. Unable to stand. 5:03. Convulsions followed by death. Experiment.—0.015 gram dried venom dissolved in 1 c. ¢. distilled water and boiled in a test-tube for about two minutes. 3:51. Injected into the breast muscles of a pigeon. 4:15. Unable to stand. 4:22. Dead. No local effects. From these experiments it appears that the toxicity of venom is not impaired by brief heating as high as 79.5° C., the time of death being in these experiments about the same as with the unheated solution. In the last two experiments in which the solution was boiled, the time of death is delayed, especially so in the last experiment, but here it must be observed that but one-half the dose was used." In one experiment made on the venom of the Copperhead (Ancistrodon contor- trix) the effect seemed to be in degree between that of the Crot us and Ancistrodon (piscivorus. Experiment.—0.03 gram dried venom was dissolved in 1 c. c. distilled water and boiled in a test-tube for two minutes. 5:00. Injected into the breast muscles of a pigeon. 5:10. Unable to stand. 5:20. Incoérdination. 6:00. Very weak. Following morning. Dead. There were very slight local effects; the blood was clotted in soft black clots; heart arrested in systole, auricles full of clots. The interior of the thoracic cavity had a mucky brownish appearance ; the viscera did not appear congested, and there were no ecchymoses. A similar dose of the unheated copperhead venom kills promptly with decided local effects. It will thus be apparent that boiling decidedly alters its toxic power. The effect of boiling on the venom of the Crotalophorus is as decided as on that of the Crotalus. Experiment.—Two drops of the fresh venom of the Orotalophorus was dissolved in 1 c.c. distilled water and boiled for a moment. 4:58. Injected into the breast muscles of a pigeon. 6:15. In good condition; no symptoms up to this time, excepting a little tendency to droop. Following evening. Animal normal. The venom of the Coral snake (Elaps fulvius) is affected to a less degree. 1 Very prolonged boiling, as has been shown by Fayrer and by Ward, lessens greatly, and at last destroys toxicity in cobra venom. The efficient cobra peptone is, as we have seen, converted into a coagulable albuminoid, which is then incapable of destroying life. 4 April, 1886. 26 THE VENOMS OF CERTAIN THANATOPHIDESA. Experiment.—Boiled 0.015 gram dried Coral venom dissolved in 1 c.c. distilled water. ; Time of injection ? 5:45. Very weak. 6:00. Nearly dead. 6:10. Dead. No local effects. Blood coagulates perfectly. A smaller amount of venom unboiled kills in from 10-15 minutes with decided local effects. From the experiments with the venom of the Crotalus adamanteus detailed above it appears as though the toxicity of the venom was completely destroyed by boiling, but Weir Mitchell found some years ago that boiling did not destroy the poison- ousness of the venom of the Crotalus durissus, and further work of our own led us to believe that the want of toxicity of our boiled solutions was only apparent, and that there was accordingly a poisonous principle still present, but not in deadly quantities. We therefore made some further observations, using larger amounts of venom. Experiment.—Dissolved three drops of fresh Crotalus adamanteus venom in 1.5 c. c. distilled water and boiled. 4:40. Injected into the breast muscles of a pigeon. 6:10. No positive effects. Following morning. Dead, no characteristic local effects. Experiment.—Dissolved 0.12 gram venom (Crotalus adamanteus) in 2 ¢. ¢. dis- tilled water and boiled for two or three minutes. 4:40. Injected the above into two pigeons, giving each half. Death within fourteen hours in both pigeons. There was some slight local effect, but nothing comparable to what is observed in the unheated venom. There were no extravasa- tions, and the blood was clotted. The stench from putrefaction at the points of injection was very great, and the muscles around them presented a pale-grayish color as though they had been boiled. A like result was obtained in the case of another pigeon experimented on in the same way. From the above series of experiments it is perfectly clear that heating the dis- solved venom beyond a definite point, varying no doubt in different venoms, lessens its toxic power. Boiling for some minutes does not destroy the poisonous capacity of the venoms, but simply impairs this quality to a varying degree, depending upon peculiarities in the toxic constituents, as we shall hereafter have reason to observe. Fayrer and Wall, as already noted, found that prolonged boiling of solutions of Cobra venom completely destroyed the poisonous activity of that secretion. We accordingly made some similar experiments with solutions of the venom of the Crotalus adamanteus with analogous results. Experiment.—0.03 gram of the dried venom of the Crotalus adamanteus was dissolved in a little distilled water and boiled for ten minutes in a water bath After being allowed to cool it was injected into the breast of a pigeon. 1:56. Injection practised. 1:57. Weak. 2:00. Convulsions. 2:37. Since last observation has been lying on its side, very weak. 2:43. Dead. EFFECTS OF VARIOUS AGENTS ON VENOM. ii In a subsequent experiment the solution of venom was boiled for forty minutes. Three minutes after the injection the pigeon vomited; no other toxic symptoms were observed. In another experiment, in which the venom was boiled for one hour, no symptoms occurred but vomiting. Both of these pigeons were watched for three days, but in neither of them did any poisonous symptoms ensue. The Effects of Alcohol.—When alcohol is added to fresh venom or to an aqueous solution of venom a copious white precipitate occurs. ‘The following experiments were made to determine if the active principles were entirely precipitated by the alcohol, and if the precipitate was poisonous. Experiment.—Four drops of the venom of the Crotalus adamanteus were placed in 1 c. c. absolute alcohol. ‘The precipitate was filtered and washed with an addi- tional amount of alcohol, the filtrate then being evaporated spontaneously to 1 ¢. ¢. The precipitate was placed in 1 c. c. distilled water and injected into the breast muscles of a pigeon at 5:11. 5:17. Too weak to stand. 5:21. Dead. There was very little local effect. The filtrate was injected into another pigeon, as above, at 5:22. € 5:26. Vomits; no further effects. From this experiment it is obvious that the presence of alcohol does not destroy toxicity. Further observations were made to learn the effect of a more prolonged action, and it the precipitate was soluble in water. Experiment.—0.06 gram of dried Crotalus adamanteus was dissolved in 3 minims of distilled water and this was added to 3 ¢. c, absolute alcohol (Squbb’s) causing a dense precipitate. ‘The mixture was allowed to stand for three days. It was then filtered, the precipitate being several times washed with the filtrate and finally with fresh absolute alcohol. The precipitate was finally washed from the filter by distilled water, allowed to dry, then digested in distilled water for twenty-four hours, and, after being filtered, was washed with distilled water. ‘The filtrate was cloudy, and on being allowed to stand for one and a half hours cleared somewhat, there being an upper layer of clear fluid and some sediment. One-fourth of the filtrate was now injected into the breast muscles of a pigeon at 4.43. 4:54. Unable to stand. 5:10. Dead. There is exceedingly little local effect. The tissues at point of injection are suffused with blood. 5:45. Blood still fluid. To one-fourth of the filtrate one minum of acetic acid was added, which caused the mixture to become clear. 4:41. Injected into a pigeon as above. 4:52. Rocking. 4:54. Down. 5:58. Dead. There is absolutely no local effect and there is no suffusion of blood in the tissues as in the previous experiment. I8 THE VENOMS OF CERTAIN THANATOPHIDESA. To one-fourth of the filtrate a few crystals of sodic chloride were added, which rendered the solution clear. 4:49. Injected as before. 4:55. Rocking. 4:58. Down. 5:57. Dead. The local effect is intense; great blackening and infiltration of fluid blood. It will have been seen that even after subjection for three days to the action of absolute alcohol the venom has not lost its toxicity. It further appears that the addition of acetic acid or sodic chloride, while rendering the undissolved material soluble delays the time of death, and that the local effects of the poison are destroyed by the acid and intensified by the sodic chloride. The action of the acid is probably due either to a powerful local constricting action on the tissues or else to a modification of the properties of the poison. We can give no reason for the cause in the delay of death after the addition of the sodic chloride. As the animal in this observation lived longer than in the first, the increased local effect may be in this way partially accounted for, The filtrate becomes very turbid by boiling, and gives a decided precipitate with nitric acid, thus proving that the water has actually dissolved some of the precipi- tate, and consequently that the toxicity of the filtrate cannot depend merely upon the undissolved particles of precipitate carried through the filter. It is interesting to learn whether alcohol dissolves any poisonous element of the venom. In one of the above experiments the only effect following the injection of the alcohol filtrate was vomiting, but the objection may be made that the alcohol was in sufficient quantity to act as a physiological antidote to any poisonous ele- ment of the venom which it might have contained. We therefore made a further test of this matter by using Cobra venom, which is more powerful than that of the Crotalus, and using it in larger quantities. Experiment.—Dissolved 0.1 gram Cobra venom in two drops of distilled water, and digested in 2.5 c.c. absolute alcohol for about ten days. ‘The mixture was then filtered, and the filtrate evaporated spontaneously to 3 of a c.c. ‘This was injected into a pigeon without any effect. The following observations with Cobra venom are of great value as throwing light upon the different results obtained by various investigators in studying the action of alcohol on venom. In this series of experiments varying proportions of water were used to dissolve the venom. Experiment.— Dissolved 0.02 gram dry Cobra venom in three drops of distilled water, then added 1 c. c. absolute alcohol and filtered. (1.) 4:37. Injected into the breast of a pigeon the above filirate—no symptoms. (II.) 4:41. Injected the precipitate in a little water. 4:50. Dead. Experiment.— Dissolved 0.03 gram Cobra in ten drops of distilled water and added 1 c.c. absolute alcohol and filtered. (1.) 5:00. Injected the filtrate as above. 5:30. Sick. 5:53. Unable to stand; extremely feeble. 5:55. Dead. EFFECTS OF VARIOUS AGENTS ON VENOM. 29 (II.) 5:05. Injected the precipitate with water. 5:074. Dead. In the first series the results are the same as in previous experiments, but in the second series, where a much larger quantity of water was used, the filtrate caused death in fifty-five minutes, thus proving that if sufficient water be present, enough of the poison is carried with the filtrate to cause death, notwithstanding the larger amount of alcohol present and its attributed antidotal action. The Action of Absolute Alcohol upon the Dried Venom.—lIf dried venom be placed in absolute alcohol and the mixture allowed to stand for some time, even for months, it will be found that the venom undergoes no change in its poisonous activity, nor does it appear that the alcohol dissolves out any of the poisonous principles, since it is found to be innocuous after injection, and does not give any reaction for proteids. The Effect of the Caustic Alkalies on the Toxicity of Venoms. Caustic Potash.— When caustic potash is added to a solution of venom the latter becomes perfectly clear. If the quantity of salt added to the solution is below a definite limit no decided alteration in the capacity to kill is noticed, but as this quantity increases obvious results are observed, first a diminution in the activity of the poison, and at last a complete loss of toxicity. Experiment.—Dissolved 0.03 gram dried Crotalus adamanteus venom in 1 ©. ¢. of distilled water in which was previously dissolved 0.0037 gram potassic hydrate. 3:45. Injected into the breast of a pigeon. 4:48. Weak. 5:00. Unable to walk; 6:00 ditto. 6:30. Dead. Heart arrested in systole; no ecchymoses; well-marked local effect ; blood fluid at the end of sixteen hours. Experiment.—The same as above, using 0.0075 gram potassic hydrate. 3:43. Injected. 5:00. Weak. 6:00. Weaker; slight local effect. Following morning. Animal living, but weak; the local effect is well marked. Experiment.—The same, using 0.015 gram potassic hydrate. 3:47. Injected. 7:00. No symptoms up to this time. 7:30. Sickish. Following morning. Sickish; some slight local effect at point of injection. Experiment.—The same, using 0.03 gram potassic hydrate. 3:50. Injected. 7:00. No symptoms up to this time. Following morning. No symptoms; no local effects. ‘This experiment was repeated in two other pigeons with a like result. The last series of experiments prove clearly that the addition of potassic hydrate to a solution of venom, if in sufficient quantity, produces a decided effect on the activity of venom, and that if added to the venom of the Crotalus adamanteus in a quantity equal to the weight of the dried poison the lethal action is entirely . destroyed. In one experiment made with the venom of the Crotalus horridus the 30 THE VENOMS OF CERTAIN THANATOPHIDE A. same holds good, but our experiments with Cobra venom show that a larger pro- portional quantity is needed to destroy its power. Experiment.—Dissolved 0.015 gram Cobra venom in 0.5 c. ¢. distilled water, then added an equal amount of potassic hydrate. 4:05. Injected into the breast muscles of a pigeon. 4:22. Unable to stand. 4:32. Convulsions. 4:37. Dead, This experiment was repeated with a similar result. A larger proportion of the potassic hydrate was used in the following observa- tions :— Experiment.—The same as above, using 0.03 gram (double the amount) of potassic hydrate. 4:51. Injected into the breast of a pigeon. No effects. Repeated this experiment with a similar result. In one instance, however, we found that 0.06 gram potassic hydrate did not effectually counteract the poisonous activity of 0.015 gram dried Cobra. It has been suggested that the non-poisonous action of venom treated with potassic hydrate and injected hypodermically, as in the above experiments, depends upon an effect of the potassic salt on the tissues, causing a considerable delay in the absorption of the poison, and this suggestion seems strengthened by the result in a rabbit of an intravenous injection of 0.015 gram Crotalus venom with 0.06 gram potassic hydrate in 1 c. c. distilled water. The animal became very sick soon after the injection, which was given in the evening, and remained in this condition at the end of an hour, when the observation ceased. ‘The following morning it was found dead, with post-mortem appearances of the effects of venom. Also, in another animal, which was given intravenously 0.015 gram of venom with a similar amount of potassic hydrate, death occurred as promptly as with pure venom; in fact rather earlier. In another set of experiments on pigeons, we carefully neutralized the potassic hydrate before injecting. We used in all of this series sulphuric acid as the neutralizing principle, so that a harmless potassic sulphate was formed. The results of this group of experiments also go to show that the potassic hydrate prevents the absorption of the venom. Experiment.—Dissolved 0.015 gram dried venom of the Crotalus adamanteus in 1 c.c. distilled water and added 0.015 gram potassic hydrate, then carefully neutralized with acetic acid. This was injected into the breast of a pigeon, causing death in sixteen minutes. Experiment.—Dissolved 0.03 gram dried Crotalus adamanteus venom in 1 ©. ¢. distilled water and added 0.015 gram potassic hydrate, and then neutralized as above. 4:19. Injected as above. 4:55. Weak; breathing rapid. 6:20. Much weaker. Following morning. Dead. Decided local effect; blood fluid and dark. EFFECTS OF VARIOUS AGENTS ON VENOM. 3l Experiment.—The same, only using 0.0075 potassic hydrate. 4:53. Injected as before. 5:00. Weak; breathing deep. 5:10. Dying. 5:18. Dead. Slight local effect; blood fluid and dark. The records of the above experiments, which are in accord with Wall’s, show that the results after the addition of the potassic hydrate are not the same as in the series where the alkali was not neutralized, thus proving that the effect of the action of the added alkali does not remain after the latter is neutralized. In previous observations we found that solutions of venom were more or less impaired by boiling, and that this was particularly marked with the venom of the Crotalus adamanteus, 0.015 gram being rendered completely innocuous to pigeons. It was afterwards found that no coagula were formed by heating solu- tions of venom to which had been added some potassic hydrate, as in the above experiments. This led us to study the results of heating solutions of venom to which the potassic hydrate was added to learn if heat was capable of destroying or impairing toxicity without the occurrence of coagulation as a necessary event. Experiment.—Dissolved 0.015 gram of the venom of the Crotalus adamanteus in 1 ¢, c, distilled water and added 0.015 grain potassic hydrate, and subjected the solution, as in previous experiments, to a gradually increasing temperature up to 74°C. It was then injected into a pigeon. At the end of twenty-four hours there was no effect. In this experiment the temperature to which the solution of venom was sub- mitted was below the point at which serious imjairment of the poisonous power of the venom occurs, yet the amount of potassic hydrate was sufficient to destroy its action. Other experiments were made in which the quantity of potassic hydrate was not sufficient to effect this end. We found in previous experiments that 0.0037 gram potassic hydrate was not sufficient to destroy the toxicity of 0.03 gram of Crotalus adamanteus venom, although the time of the occurrence of death was considerably delayed. ; We used similar amounts of venom and alkali in the three following experi- ments, using 0.95 c. c. distilled water for the solutions. Experiment.—Dissolved 0.09 gram of Crotalus adamanteus venom in 1.5 ec. ec. distilled water and added 0.011 gram of potassic hydrate. This solution was divided into three parts. One of which was heated to 76.5° C., one to 79.5° C., and the other to 83.5° C. Each of which was injected into the breast of a pigeon and without any evil consequence following within twelve hours. These results indicate that heat impairs the poisonous activity of venom under the above conditions, even though coagulation does not occur. In previous expe- riments recorded it was found that at a temperature of 79.5° 0.03 gram of Crotalus adamanteus venom was rendered non-toxic. The explanation of the further impairment of the action of the poison by heating its solutions having potassic hydrate dissolved in them lies probably in the fact that the potassic hydrate is placed by heat under condition of greater activity. The non-coagulability of solutions of venom to which potassic hydrate was added is no doubt due to the alteration By) THE VENOMS OF CERTAIN THANATOPHIDEA. of the coagulable proteids into alkali-albumins, and as a moderate degree of heat increases the rapidity of this change, it is possible that the smaller amount of alkali is as effective under these conditions as the larger amounts under ordinary conditions. It is not at all improbable that the prolonged action of potassium hydrate on solutions of venom may convert all of the globulins into alkali-albumins and thus destroy their poisonous activity. Sodic Hydrale-—The effect of sodic hydrate on solutions of the venoms of the Crotalus adamanteus and horridus appears to be the same as that of the potassic salt. In one experiment with the Crotulus adamanteus, using equal quantities (0.03 gram) of the dried venom and alkali, no poisonous effects followed its injec- tion; and in another experiment in which 0.015 gram of venom and 0.007 gram sodic hydrate were used the animal was rendered somewhat sick, but fully recovered. In one experiment with the venom of the Crotalus horridus, using equal quan- tities (0.015 gram) of the venom and sodic hydrate, no poisonous symptoms followed. The effect on solutions of dry Cobra venom, as in the case of the potassic salt, is not so marked. Experiment.— Dissolved 0.015 gram dry Cobra yenom in 0.5 ¢. ¢, distilled water and added (0.015 gram sodic hydrate. 4:08. Injected into the breast of a pigeon. 4:15. Unable to stand. 4:27. Dead. In two other experiments, using double the quantity of sodic hydrate, one animal died in one hour, and the other in a little less than three hours. Double amounts therefore decidedly impair toxicity, In another experiment, in which four times the quantity of sodic hydrate was used (0.015 gram dried venom + 0.06 gram NaHO), no poisonous symptoms followed.’ The Effects of Ammonia.—The dry venom of the Crotalus adamanteus, which was the only. one used, forms with aqua ammonia a turbid solution, such as is formed with water. The effect on the toxicity of the venom exerted by the ammonia is not so marked as with the potassic or sodic hydrates. Experiment.—Dissolved 0.08 gram dried venom in two minims aqua ammonia (20°) with 1 ¢. c. distilled water. 5:29. Injected into the breast muscles of a pigeon. 5:37. Unable to walk. 5:46. Convulsions; death. The local lesions are decidedly lessened by the alkali. Experiment.—The same, using six minims aqua ammonia. 4:14. Injected as above. 6:00. No marked symptoms up to this time, excepting droopiness. The local effect is slightly more marked than in No. 1. Following morning the animal was dead. In three other experiments in which eight minims of aqua ammonia were used two of the animals were found dead the following morning and one recovered. In * See Shortt. Wall, op. cit., p. 133. On the effects of alkalies and of permanganates, see Vincent Richards, F.R.C.S. Ed., ete., op. cit. EFFECTS OF VARIOUS AGENTS ON VENOM. 33 another experiment in which the alkali was neutralized by sulphuric acid, death did not occur for four hours. In the first experiment, in which a very small amount of ammonia was used, death occurred in less than twenty minutes; in the next, in which three times the quantity of ammonia was used, death did not ensue for some hours, while in the next three a more positive effect was no doubt apparent in the fact that one of the pigeons recovered. In the last experiment death did not occur for over four hours, even after neutralization of the alkali, indicating, as in the case of the potassic hydrate, that some permanent effect had been exerted on the venom by the ammonia. Potassium Carbonate.—Two experiments made with. the venom of the Crotalus adamanteus render it probable that the potassic carbonate does not exert any decided effect. Experiment.—Dissolved 0.015 gram venom in 1 ¢.c. distilled water and added 0.015 gram potassic carbonate. 4:16. Injected into the breast of a pigeon. 4:22. Down. 4:25. Dead. No appreciable local effect. Experiment.— Dissolved 0.03 gram venom in 1 c.c. distilled water and added 0.12 gram potassic carbonate. 5:25. Injected into the breast muscles of a pigeon. 5:45. Down; observation now ceased. Following morning found dead; slight local effect. Nitric Acid.—The powerful destructive action exerted by this acid on albumi- noids suggests at once that it would in all likelihood completely destroy the poison- ous properties of venom, yet it has been asserted that such is.not the case. In the latter instance the result was no doubt due to the insufficiency of acid used, as we have clearly determined in our experiments. Experiment.—Dissolved 0.03 gram Crotalus adamanteus venom in 0.5 c. c. dis- tilled water and added 24 minims C. P. nitric acid, which caused a considerable precipitate. 3:32. Injected the above into the breast of a pigeon. 3:33. Convulsions, followed by death. From this result it seemed probable that not enough acid had been added to throw down all of the precipitable proteids. In another experiment the acid was added to a solution of venom and the mixture filtered. he filtrate was now tested with nitric acid and a further precipitate occurred. This process was repeated until no further precipitate followed. The filtrate was set aside, and the precipitate on the filter washed with dilute nitric acid and then with water. Experiment.—5:05 injected into the breast of a pigeon the above filtrate, which measured 3 c. c. and contained 1 ¢. ¢, nitric acid. 6:05. No symptoms except slight droopiness. Following morning no effects from venom. 5 @April, 1886. ot THE VENOMS OF CERTAIN THANATOPHIDES. Eaperiment.—5:55 injected the precipitate in 1 ¢, c, dilute nitric acid with which it had been in contact for two hours. 6:05. No symptoms. Following morning animal in good condition. It will thus be observed that the acid has completely destroyed the toxicity of venom. We made still another experiment in which the venom was rubbed up in a mortar and the acid added to it, and then diluted with water. Experiment.—0.03 gram dried Crotalus venom was rubbed in a mortar until powdered, and 4 gtt. C. P. nitric acid added. ‘This formed a pasty mass of an orange-yellow color. With 1 ¢.c. distilled water it formed a cloudy, orange-_ yellow solution. The above was injected into the flank of a half-grown rabbit, without any symp- toms of venom poisoning following within twelve hours. A similar experiment was made with a pigeon with a like result. The acid, however, having been neutralized with sodic carbonate before injection. Muriatic Acid.—This acid does not seem to exert so strong an effect. Only one experiment was made. Experiment.—0.015 gram dried Crotalus venom was rubbed in a mortar, and to it was added 4 gtt. C. P. muriatic acid forming a clear solution. With 1 c.c. distilled water it made a turbid solution. ' 8:44. Injected the above into the breast muscles of a pigeon. 5:00. Very sick. 5:50. Nearly dead. Following morning dead; no local lesions from venom. Here the amount of venom used was only one-half of that employed in the nitric acid experiment. ‘The quantity of acid was the same, but in this experiment a pigeon was used. As in the series with nitric acid, an experiment was also made in which the dried venom was powdered in a mortar and a few drops of the pure acid used. About 1 c.c. of distilled water was added, and the mixture neutralized with sodic carbonate. It was then injected into the breast of a pigeon with the result of death in twenty-six minutes. Sulphuric Acid.—Repeated the above, using instead of the muriatic acid 5 ett. sulphuric acid. The venom and acid formed a clear syrupy solution which became milky by the addition of the water. . 8:53. Injected as above. 5:50. Sickish. Following morning dead; no local symptoms of venom poisoning. Dr. Mitchell had observed that if the acid was afterwards neutralized the action of the venom was not affected. ‘The delay of death in this experiment seems to be due to the action of the non-neutralized acid. We, however, made an experiment by powdering the dried venom (0.015 gram) in a mortar, adding a few drops of the pure acid, diluting then with about 1 c. c. distilled water, and neutral- izing with sodic carbonate. ‘This was injected into the breast of a pigeon. HFFECTS OF VARIOUS AGENTS ON VENOM. 35 For some time after the injection the bird was weak, and continued in a feeble condition until eighteen hours after the injection, when death ensued. It seems quite remarkable that such a powerful acid as sulphuric does not com- pletely destroy the poisonous properties of the venom, and it is even more curious that pure muriatic acid seems to be without effect. Acetic Acid. Experiment.—Dissolved 0.02 dried venom (Crotalus adamanteus) in 0.1 c. c. distilled water and added 3 minims of glacial acetic acid. 4:30. Injected into the breast of a pigeon. 4:37. Incodrdination. 4:41. Dead. Death occurred in this experiment in such a short time that it was thought that the acid itself might have contributed to this end. We therefore made another experiment m which the acid was neutralized. Experiment.—Prepared the venom as before, only neutralizing the solution with sodic carbonate. 4:35. Injected into the breast of a pigeon. 5:10. Pigeon unable to stand. 5:15. Dead. The result of this experiment indicates that the presence of the free acid aids the toxic action of venom. Hydrobromic Acid. Experiment.—Powdered 0.015 gram dried Crotalus ada- manteus venom in a mortar and added 5 gtt. hydrobromic acid (sp. gr. 1.274), after five minutes added 0.5 c. c. distilled water. ‘The venom and acid formed a slightly reddish-colored solution, which became milky when diluted with water. 4:25. Injected into the breast muscles of a pigeon. 4:45. Sickish. 4:55. Unable to stand. (Final result: not noted, but death most certainly followed.) -We repeated the above experiment, using 10 gtt. of acid mixed with an equal part of water, before dissolving the venom in it. 2:49. Injected as above. 3:07. Rocking. 3:30. Dead; local effects of the venom apparent. Notwithstanding we used double the amount of acid in this experiment, it does not appear as though the activity of the venom was made to differ much from that noted in the previous experiment. Since the previous dilution of the acid before mixing with the venom might have affected its action a third experiment was made in which the same quantity of acid was added, without dilution, to the powdered venom. Hxperiment.— Powdered 0.015 gram dried venom and added 10 gtt. hydro- bromic acid, then 1 ¢. c. distilled water. 4:20. Injected the above into the breast muscles of a pigeon. 5:00. No apparent effect. 5:10. Sickish. 6:00. Sickish. Following evening. Well. 36 THE VENOMS OF CERTAIN THANATOPHIDESA. This last experiment was repeated with the modification of leaving the acid in contact with the venom for one-half hour before the addition of the water. It was then injected as above without any obvious effects following. The destructive action of the acid on the venom of the Crotalus horridus seems to be the same if we can judge from the single experiment which follows. Experiment.—Powdered 0.015 gram dried venom and added 10 gtt. hydro- bromic acid, which formed a muddy solution with a reddish color. 5:18. Injected into the breast of a pigeon without any obvious effects within twenty-four hours. The effect on the activity of Cobra venom, using similar quantities of venom and acid is very different. Experiment.—Repeated the above, only substituting Cobra venom. 4:48. Injected into the breast muscles of a pigeon. 5:18. Sick; breathing difficult. 5:30. Breathing more difficult; convulsive movements; incodrdination. 5:35. Dead. Tannic Acid.—The action of tannic acid upon albuminoids is so decided that we might confidently expect, since we find the poisonous elements in venoms to be proteids, that the activity of venom would be greatly diminished or entirely destroyed by it. In one experiment made with the venom of the Crotalus adu- manteus we found comparatively little effect. Experiment.—Dissolved 0.03 gram dried venom in a little distilled water and added 1.5 c. c. saturated solution of tannic acid. 3:35. Injected into the breast muscles of a pigeon. 4:00. Droopy. 4:45. The same. Following morning dead. It will be observed that there is a great delay in the action of the venom, possi- bly due to the powerful local constrictive action of the tannic acid on the tissues, and possibly, also, to a direct action of the acid on the venom itself. As death may have resulted from the tannic acid we made a control experiment in which 1.5 c. c. saturated solution was injected into the breast of a pigeon. ‘The animal did not exhibit any signs of active poisoning, but it died at the end of the fourth day. Alum.—We made but two experiments with alum, one with the venom of the Crotalus horridus and one with Cobra. Experiment.—Dissolved 0.015 gram dried venom in 0.5 c. c. distilled water and added 3 gtt. saturated solution of alum (18° C.), but no precipitate occurred; we then gradually added powdered alum nearly to saturation, which caused precipita- tion. The precipitate was filtered off, and the clear filtrate tested by the further addition of alum to see if any more precipitation would occur, with a negative result. The precipitate and filtrate were now mixed together and injected into the breast of a pigeon without any poisonous result occurring within forty-eight hours. In another experiment in which 0.06 gram of dried Crotalus venom was used, the animal died in forty-five minutes. EFFECTS OF VARIOUS AGENTS ON VENOM. By7/ Alum added to saturation does not precipitate the peptone, although it precipi- tates all of the coagulable proteids. The following is the experiment with Cobra venom:— Experiment.—Dissolved 0.015 gram dried venom in 0.5 c. c. distilled water and added alum to saturation (16° C.). 4:32. Injected into the breast muscles of a pigeon. 4:50. Down. 4:52. Dead. This last experiment is of interest in proving that even so powerful an astringent as alum is not sufficiently strong to prevent the prompt absorption of the poison. Death followed in twenty minutes. Chlorine Water.—This reagent does not seem to exert any influence. Experiment.—Dissolved 0.015 gram Crotalus adamanteus venom in 0.5 ¢. ¢. dis- tilled water and added 0.5 c. c. fresh chlorine water. 4:28. Injected into the breast muscles of a pigeon. 4:52. Down. 5:10. Dead. Bromine.—The action of bromine in bromohydric acid solution is very marked. Experiment.—Powdered 0.015 gram dried Crotalus adamanteus venom in a mortar and added 2 gtt. of bromine in 4 or d gtt. bromohydric acid, then added 0.5 c. c, alcohol. 5:05. Injected into the breast of a pigeon. 5:30. No effect. Twenty-four hours. No effect. This experiment was repeated once with Crotalus venom and once with Cobra, using water as the diluent instead of alcohol. In both experiments we found a similar result, thus proving that the activity of the venom is completely destroyed by this reagent. Jodine. Eaxperiment.—Dissolved 0.015 gram dried venom of Crotalus ada- manteus in 0.33 c. c. distilled water, then added 0.5 c. c. tr. iodine which formed a dense brown precipitate. 5:07. Injeeted into the breast of a pigeon. No poisonous effects within twenty-four hours. If, however, the amount of iodine be much smaller the venom is still potent, as is shown by the following experiment. Experiment.—Dissolved the venom as above, then added 1 drop tr. iodine and afterwards 1 c.c. distilled water. 4:56. Injected into the breast of a pigeon. 5:05. Weak. 5:15. Dying. Todine 4- Potassic lodide. Experiment.—Dissolved 0.015 gram dried venom of Crotalus adamanteus in 0.5 ¢. c. distilled water, then added a saturated solution of equal parts of tr. iodine and potassic iodide. 4:41. Injected the above into the flank of a small rabbit (half grown). The animal died in about eighteen hours. 38 THE VENOMS OF CERTAIN THANATOPHIDESA. The delay in the occurrence of death in this experiment was considerable, and that this was due to the action of the iodine on the venom is rendered probable by the results of the preceding experiments with iodine and by the following experiment with the potassic iodide. Potassic Iodide.—This salt does not seem to exert any influence upon the activity of venom. Experiment.—Dissolved 0.015 gram dried venom of the Crotalus adamanteus in 1c. c. saturated solution of potassic iodine. 4:31. Injected into the breast of a pigeon. 4:40. Down. 4:45. Dead. Potassic Bichromate. Experiment.—Dissolved 0.03 gram dried Crotalus ada- manteus venom in 1 ¢. ¢. distilled water and added 0.01 gram potassic bichromate. 4:14. Injected into the breast of a pigeon. 4:20. Down. 4:25. Convulsions followed by death. Experiment.—Dissolved 0.004 gram dried venom in 0.5 c. c. distilled water and added 0.03 gram potassic bichromate dissolved in 0.33 distilled water, which pro- duced a dense coagulum. 3:38. Injected into the breast muscles of a pigeon. 4:05. Dead. Potassic Permanganate. Experiment.—Dissolved 0.03 gram dried Crotalus adamanteus venom in ().5 c, ¢, distilled water and added 0.06 gram permanganate in 0.5 ¢. c. distilled water. ‘This formed a very cloudy solution. 5:27. Injected into the breast of a pigeon. Death occurred within forty-eight hours. Experiment.—The same, using 0.015 gram of the permanganate. At the end of the second day no poisonous effects from the venom. ‘The parts where the injec- tion was made look as though they would slough. Experiment.—The same, using 0.005 gram of the permanganate. The solution formed is a dark wine color. 4:37. Injected into the breast of a pigeon without effect. Eaxperiment.—The same, using 0.0038 gram of permanganate. 4:26. Injected as above. 4:42. Down. 4:45. Dead. Experiment.—The same, using 0.0025 gram of permanganate. 3:57. Injected as above. 4:06. Down. 4:10. Dead. In another experiment, the mixture was injected into the femoral vein of a rabbit, using 0.005 gram permanganate. The animal lived, and at the end of the second day was apparently unaffected. In one observation made with the venom of the Crotalus horridus 0.015 gram of venom was dissolved in 0.5 c. c. distilled water, to which was afterwards added 0.008 gram of the permanganate. After standing for twenty-four hours the EFFECTS OF VARIOUS AGENTS ON VENOM. 39 mixture was very thick and tarry, and would not flow from the inverted test-tube. It seems from this that the full extent of the action of the permanganate on the venom is not exerted for some hours. The permanganate is efficient in destroying the activity of Cobra venom. Experiment.— Dissolved 0.015 gram dried Cobra venom in 0.5 c. ¢. distilled water and added 0.015 gram permanganate. 4:35. Injected into the breast muscles of a pigeon. No symptoms of venom poisoning within twenty-four hours. ‘ Peroxide of Hydrogen. —Notwithstanding the powerful nature of the peroxide of hydrogen as an oxidizer, it does not seem to affect to any great extent the poisonous activity of venom. Only one experiment was made. Experiment.—Added 3 drops of fresh venom of the Crotalus adamanteus to 3 c. c. fresh solution of peroxide of hydrogen, specially prepared by Prof. Leeds, of Hoboken. 9:05. Injected into the breast muscles of a pigeon. 5:15. Unable to stand; decided local effects appearing. 6:05. Dead, with all the usual phenomena of venom poisoning. The quantity of peroxide of hydrogen used in this experiment was so large that the test was a satisfactory one. Argentic Nitrate.— Notwithstanding the powerful action of nitrate of silver on albuminoids it does not seem to possess great power to disturb the toxicity of venom. Experiment.— Dissolved 0.015 gram of dried venom of Crotalus adamanteus in 3 ¢.¢. distilled water, to which was afterwards added 0.015 gram nitrate of silver, forming a decidedly milky solution. 4:40. Injected into the breast of a pigeon. 4:50. Down; deep breathing, gasping. 4:53. Dead. : As there was a possibility of the quantity of salt being insufficient for the amount of venom, another experiment was made in which double the weight of nitrate was used. The mixture was injected into the breast of a pigeon. At the end of three days no symptoms of venom poisoning had occurred. Mercurie Chloride-—When mercuric chloride is added to a solution of Crotalus or Moccasin venom a dense precipitate occurs, consisting of all the proteids in solution. In order to learn if the precipitated proteids still retained any toxic power we dissolved 0.03 gram of dried venom of the Crotalus adamanteus in 1 ce. c. distilled water and then added 0.03 gram mercuric chloride. The precipitate was collected on a filter and repeatedly washed with distilled water. During this washing the precipitate seemed to diminish a little in quantity, and was no doubt partially dissolved. 3:30. The precipitate in 1 ¢. ec. distilled water was injected into the breast of a pigeon. 6:00. No symptoms up to this time. Twenty-four hours—the animal showed no signs of venom poisoning. Ferrous Sulphate——Three experiments were made with the sulphate of iron with results materially different; the difference no doubt depending upon the mode 40 THE VENOMS OF CERTAIN THANATOPHIDEA. of administration. In all the same quantities of venom and salt were used, but in one the solution was injected simply beneath the skin and in the others directly into the muscles of the breast. In the former the animal did not die until after the lapse of nearly thirty-six hours, while one of the others died remarkably soon —within four minutes after the injection, and the third in twenty-eight minutes. Experiment.—Dissolved 0.03 gram dried venom of the Crotalus adamanteus in 1 c. c. distilled water and then added 0.03 gram ferrous sulphate. The addition of the iron salt renders the solution clear. 3:40. Injected beneath the skin of the thigh of a pigeon. 6:00. No apparent effects. Twenty-four hours—no effects. Thirty-six hours—dead. Slight local effects of venom, but the destructive action of the iron salt on the tissues is much more prominent. Experiment.—The same as above. 3:32. Injected into the breast muscles of a pigeon. 3:36. Convulsions; death. No local lesions. In another experiment the bird died in twenty-eight minutes after injection. It must be concluded from this that the ferrous sulphate does not destroy the activity of the venom. Dialyzed Iron.—When dialyzed iron is added to a solution of venom all of the proteid matter is precipitated, and the filtrate is found to give no reaction for proteids with the xanthoproteic or picric-acid tests. The precipitate is brown, and so gelatinous that if the solutions are somewhat concentrated it does not flow. The precipitate does not dissolve in distilled water, yet it must be very soluble in the tissues since the toxic effects of the venom rapidly appear after its injection. We made two experiments, both with Moccasin venom, one with the dried and the other with fresh venom. Experiment.—Dissolved 0.015 gram dried Moccasin venom in 0.5 ce. c. distilled water and added 3 gtt. dialyzed iron. ‘This caused a considerable amount of brownish gelatinous precipitate which thickened the mixture appreciably. Now added 1 c. c. distilled water. 3:20. Injected into the breast muscles of a pigeon. 3:25. Down. 3:45. Dead. Experiment.—Took two drops of fresh Moccasin venom and added first 5 gtt. dialyzed iron, and then 1 c. c. distilled water. The iron and venom made a very thick brownish mixture. 5:18. Injected into the breast muscles of a pigeon. 5:30. Dead. One experiment was made in this connection to see if dialyzed iron exerted any poisonous effect, we injected thirty drops into the breast muscles of a pigeon, without toxic result. Ferric Chloridé.—We have used the chloride of iron in two forms; the officinal tincture, U.S. P., and the officinal liquor. Both these solutions greatly affect the poisonous activity of venom, the latter, indeed, if used in sufficient quantity, EFFECTS OF VARIOUS AGENTS ON VENOM. 41 wholly prevents the occurrence of any of the symptoms of venom poisoning. The tincture does not appear to be nearly as efficient. Experiment.—Dissolved 0.015 gram dried venom of the Crotalus adamanteus in 0.5 c.c. distilled water and added 10 gtt. ¢r. chloride of iron. As the iron was added the solution cleared, but in a few moments became milky, and finally thick with whitish precipitate. 4:22. Injected into the breast of a pigeon. 5:00. No symptoms. 5:45. No symptoms. Following morning—dead. No local effect. In two similar experiments, in which double the quantity of the tincture of iron was used, the result was much the same, the time of death being notably delayed. The following experiments were made with the liquor :— Eaperiment.—Dissolved 0.015 gram dried venom of Crotalus adamanteus m 0.5 c. c. distilled water and added 4 gtt. liquor ferri chloridi. A heavy precipitate fell. 4:45. Injected into the breast of a pigeon. 5:00. Very quiet. 6:00. No symptoms, and none of venom poisoning within two days. A similar experiment was made with identical results. In one experiment with the venom of the Crotalus horridus, in which only two drops of the liquor were used, the animal showed no evidences of poisoning; and in four experiments made with the dried venom of the Moccasin, in which 0.015 gram of dried venom was used and eight, four, two, and one drop of the liquor were used, three animals gave no symptoms of venom poisoning, and one died on the third day—the animal receiving the injection containing but one drop of the iron, ‘This was the only pigeon of the four which gave any signs of poisoning. In three-fourths of an hour the bird was shaky, and at the end of three hours decidedly feeble, remaining pretty much in this condition until death. About the point of injection the iron produced considerable hardening of the tissues. The effect on Cobra venom is not marked, although in one experiment there was an appreciable delay in the occurrence of death; but in the other, in which the quantity of iron was larger, death occurred with remarkable rapidity. Ezperiment.—Dissolved 0.015 gram dried Cobra venom in 0.5 c. c. distilled water and added 2 drops liquor ferri chlor. A slight precipitate occurred in the solution after a few moments. 3:47. Injected into the breast muscles of a pigeon. 4:15. Convulsions. 4:27. Dead. Eaperiment.—Dissolved 0.015 gram dried Cobra venom in 1 c. c. distilled water and added 5 gtt. sol. perchloride of iron. This was injected into the breast of a pigeon, with the result of death in twenty seconds. The reason why ferric chloride is inefficient in destroying the toxicity of Cobra venom no doubt lies in the fact of its main poisonous substance being a peptone, 6 April, 1886. 42 THE VENOMS OF CERTAIN THANATOPHIDEA. and, like that element in all the venoms, unaffected by the iron, while the principal toxic effects of the Crotalus and Ancistrodon venoms is due to the globulins which are precipitated and chemically altered by the iron salt. Filiration through Various Substances.—Filtration through alumina or wood charcoal does not affect the poisonous activity of the venom, but by filtration through animal charcoal all of the poisonous material in venom is left behind and the filtrate is accordingly innocuous. Experiment.—Dissolved 0.03 gram dried Moccasin venom in 2 e. c. distilled water and filtered four times through animal charcoal. ‘The filtrate gives no proteid reaction. 4:20. Injected 1.5 c. c. of the filtrate into the breast of a pigeon. At the end of twenty- four hours no symptoms of venom poisoning had occurred, but there was some cedema at the point of injection. Repeated the above experiment, using 0.045 gram of Moccasin venom, and with similar results. Snake Bile—Among the curious substances which have been extolled as anti- dotes for venom poisoning is snake bile. We made but one experiment, which speaks volumes. Experiment.— Mixed 14 minims of fresh venom from a dead Crotalus adam- anteus with 1 c. c. of bile from the same animal. 4:47. Injected into the breast of a pigeon. 4:474. Incoodrdination. 4:50. Gasping respiration. 4:55. Convulsive movements. 4:56. Dead. Digestion. By digestion in strong artificial gastric juice made from the pig’s stomach the toxic power of venom (Crotalus) is completely destroyed. Experiment.—Vhree drops of the glycerine solution of venom (Crotalus horridus) (1862) were digested for sixteen hours in about 1 ¢. c. fresh artificial gastric juice from the pig’s stomach. 8:30 A.M. Injected into the breast muscles of a pigeon. Up to the end of forty-eight hours no poisonous symptoms ensued. This experiment was repeated with an identical result. We also made two experiments in which the digestive process was not carried on for such a length of time, in both only four hours, and with similar results. In one we used six drops of the glycerine solution of venom as above—just double the dose—and in the other 0.015 gram of the dried Crotalus adamanteus venom. The results of digestion in artificial pancreatic juice are similar. We made but one experiment, and that with the venom of the Crotalus adamanteus. Experiment.—Digested 0.03 gram dried venom in 1 c. c. freshly prepared pan- creatic juice from the pig for twenty-four hours. 3:44. Injected into the breast muscles of a pigeon. 5:45. Slightly droopy. Following morning, no effects apparent. EFFECTS OF VARIOUS AGENTS ON VENOM. 43 Résumé.—The above experiments, with others too numerous for detail, have enabled us to confirm Lacerda’s and Vincent Richard’s views as to the power of permanganate of potassium to destroy venoms. As a local antidote it is for all snake poisons the best. It is also clear from what we have seen that ferric chloride is a very efficient local destroyer of the venom of our own snakes, which owe their vigor to venom- globulin, but has little value as a local antidote to the peptone which gives power to the poison of the Cobra. ‘he chloride needs to be locally used in full doses, whence it is that the strong liquor ferri chloridi (U.S. P.) is more efficient than the tincture. That bromine may prove valuable as a local means of relief seems to be plain from our experiments, and is in fact one of their most interesting results. It was used, as we have seen, either as hydrobromic or bromohydric acid. Probably any solution of bromine would answer, and—as was shown by its free local use to control gangrene during our civil war—there need be no fear in using it with freedom. It has long been known in India that the strong alkalies destroy venom, and this we are able to confirm, Brainerd long ago taught that iodine has destructive value as regards Crotalus venom, and this also seems to us to be true. In fact many agents more or less alter venoms if allowed to remain long in con- tact with them, and usually act with increased vigor as the temperature is raised above that of the air; but it is chemically singular that brief exposure of venoms to strong acids should so little affect the toxicity of the poisons in question. Except where otherwise distinctly stated, the chemicals used by us have been added to the poison immediately before injecting it. Enough has been here proved to make it now worth while to study still more carefully the value of bromine and ferric chloride as local poison destroyers. One agent may be at hand or available when others are not, and the more numerous are the means we possess as local antidotes the better is the chance of escape or relief for persons bitten. 44 THE VENOMS OF CERTAIN THANATOPHIDESA. CHAPTER IV. THE EFFECTS OF VENOM WHEN APPLIED TO MUCOUS OR SEROUS SURFACES. The Effects of Venom when applied to Mucous Surfaces.—The question of the absorption of venom by mucous surfaces is one of great interest, and the verdict of all observers in connection with the venom of the Crotalide is that uninjured mucous surfaces, except in the lungs, cannot absorb venom, at least in sufficient quantities to produce death. In experiments with the venom of the Cobra other investigators have gotten results which are directly contrary, but in our own researches a large proportion of the animals survived. In seven experiments made on pigeons, in which a solution of Cobra venom was placed in the craw by means of an cesophageal tube, six showed no evidences of poisoning and one died. In these experiments the cesophageal tube, which was a small catheter, was oiled and passed with great care into the crop, the solution of venom was then poured through the tube by means of a funnel, and afterwards washed down with a little water. Experiment.—Dissolved 0.025 gram dried Cobra (Naja trip.) m about 1 e. ¢. distilled water, and placed it in the crop of a pigeon by means of an cesophageal tube. Up to four days the animal showed no signs of poisoning. Five other experiments, like the above, gave identical results. In one experi- ment the animal died within twelve hours. Experiment.—Dissolved 0.013 gram dried Cobra (Naja trip.) in about 1 e. ¢. distilled water and gave to a pigeon, as above, at 4:00 p.m. tO eH 10 30 00 30 00 30 @) 2) 00 30 30 Time: min. sec. Com Fe DP ee Time: min. sec. 20 30 40 50 0.0 1 20 3y 00 5 50 6 00 8 00 10 00 12 00 14 00 19 00 1 1@) ORS2.0 1g) 19 40 19 50 00 Pulsations per minute. 295 300 300 300 295 300 305 290 285 Pulsations per minute. 315 320 33 330 320 340 340 350 Pulsations per minute. 240 300 300 300 300 bo bo bo bo feo pO bO tO Pb OATaTANOSaTH DP Ke Hee ne Se Se , REMARKS. Pneumogastric nerves cut. Injected intravenously 0.003 gram dried venom of the Crotalus adamanteus dissolved in 1 c. ec. distilled water. Injected a similar dose. Dead. REMARKS. Pneumogastric nerves cut. Injected intravenously 0.015 gram dried venom of the Crotalus horridus dissolved in lc.c. distilled water. Violent struggles. Dead. REMARKS. Pneumogastric nerves cut. Injected intravenously 0.003 gram dried venom of the Ancistrodon piscivorus dissolved in 1 ¢.c. distilled water. Struggles. Struggles. Convulsions. Injected a similar dose. THE ACTION OF VENOMS UPON THE PULSE-RATE. 65 Time: Pulsations REMARKS, min. sec. per minute. 20 20 240 21 20 255 23 20 300 25 20 270 28 20 255 33 «20 255 38 20 259 44 20 255 Injected a similar dose. 44 40 225 45 10 soo, Den. Experiment No. 24. Time: Pulsations REMARKS. min. sec. per minute. Normal Nea 300 Pneumogastric nerves cut. Injected intravenously 0.003 10 300 gram dried venom of the Ancistrodon contortrix dissolved 20 310 in 1c. e. distilled water. 30 310 : 40 310 1 00 310 i XO) 310 4 20 310 7 00 310 Injected a similar dose. T 05 ao Struggles. i 10 320 eee) 300 7 40 300 8 00 310 9 00 315 it BO 300 Injected a similar dose. 11 40 295 12 00 290 12 3 290 13 00 290 13 30 290 19 00 285 - 19 20 270 19 40 270 20 20 285 21 50 285 22 50 285 Experiment No. 25. - Time: Pulsations REMARKS. min. sec. per minute. Normal oo 6 330 Pneumogastric nerves cut. Injected intravenously 0.003 10 330 gram dried Cobra venom dissolved in 1 e. e. distilled water 30 330 with a few crystals of sodic chloride and filtered. 0) 330 3 30 340 6 30 330 10 630 , 320 14 30 295 16 30 275 Clot formed in canula. 9 May, 1886. 66 THE VENOMS OF CERTAIN THANATOPHIDEA. Experiment No. 26. Time: Pulsations — REMARKS. min. sec. per minute. Normal Bo 8 320 Pneumogastrie nerves cut. Injected intravenously 0.006 10 315 gram dried Cobra venom prepared as in the foregoing 30 330 experiment. ‘ i OO 330 1 30 330 3 3 335 5 30 340 8 30 320 11 30 330 18 30 Bea Convulsions; asphyxia; death in 24 minutes. Experiment No, 27. Time: Pulsations REMARKS, min. sec. per minute. Normal Papi: 390 Pneumogastric nerves cut. Injected intravenously 0.003 0 5.00 gram dried Cobra venom dissolved in 1 e. c. distilled water. 10 nhs 20 396 10,0 390 2 00 360 4 00 354 10 Clot in canula. 15 Dead of asphyxia. The Actions of Pure Venoms on Animals in which Sections of the Pneumogastric Nerves and of the Upper Cervical Portion of the Spinal Cord had been made.— After isolation of the heart from the nerve centres by making section of the pheumogastric nerves and spinal cord in the middle or upper cervical region, and maintaining the animal alive by means of artificial respiration, we find that the pulsations of the heart are almost invariably slightly diminished in frequency upon use of venom. Seven experiments were made: three with the venom of the Crotalus adamanteus ; one with the Crotalus horridus ; one with the Ancistrodon piscivorus ; one with the Ancistrodon contortrix, and one with Cobra. In one expe- riment with the Crotalus adamanteus in which two doses were given, there occurred a diminution after the first dose, while there was a marked increase after the second, In the experiment with the Crotalus horrvidus there was but little alteration. Experiment No, 28. Time: Pulsations REMARKS. min. sec. per minute. Normal fee 240 Pneumogastrie nerves and cord cut. Injected intravenously 10 235 0.003 gram dried venom of the Crotalus adamanteus dis- 20 230 solved in 1 c. c. distilled water. 30 225 40 215 1 00 210 1 20 1 40 2 2 40 afore Dead. THE ACTION OF VENOMS UPON THE PULSE-RATHE. 67 Experiment No. 29. Time: Pulsations REMARKS. min. sec. per minute. Normal G corks 185 Pneumogastric nerves and cord cut. Injected intravenously 10 185 0.003 gram dried venom of the Crotalus adamanteus dis- 20 185 solved in 1 c. c. distilled water. 30 180 1 00 180 1 20 180 3 20 180 3 50 180 5 50 160 7 50 ahs Dead. Experiment No. 30. Time: Pulsations REMARKS. min. sec. per minute. Normal Balin 240 Pneumogastrie nerves and cord cut. Injected intravenously 10 230 0.003 gram dried venom of the Crotalus adamanteus dis- 20 230 solved in 1 ¢. c. distilled water. 30 230 40 195 1 00 200 20) 205 1 40 210 2 00 230 2 30 265 3 00 250 6 00 300? Injected a similar dose. 6 05 265 1165) are Gasp 270 tt 05 260 t 8 260 8 05 260 1505 O° 6 Dead. Experiment No. 31. Time: Pulsations REMARKS. min, sec. per minute, Normal ovo, 235 Pneumogastric nerves and cord cut. Injected intravenously 10 230 0.015 gram dried venom of the Crotalus horridus dis- 30 240 solved in 1c. e. distilled water. 40 240 1 00 235 2 00 240 4 00 240 6 00 230 8 00 220 12 00 ouer Dead. 68 THE VENOMS OF CERTAIN THANATOPHIDESA. Experiment No, 32. Time: Pulsations REMARKS. min. sec. per minute. Normal Suess 220 Pneumogastric nerves and cord cut. Injected intravenously 20 210 0.003 gram dried venom of the Ancistrodon piscivorus dissolved in 1 c. ec. distilled water. 30 200 Struggles. 40 200 i OO 210 1 30 210 1 60 210 4 20 195 8 20 210 10 20 210 12 20 210 15 20 210 18 20 210 21 20 Dead. Experiment No. 33. Time : Pulsations REMARKS. min. sec. per minute. Normal eae 260 Pneumogastric nerves and cord cut. Injected intravenously 10 255 0.003 gram dried venom of the Ancistrodon contortria 20 250 dissolved in 1 ¢. c. distilled water. 4() - 243 100 243 130 245 2 00 240 4 00 240, 7 00 240 9 00 240 OO 240 13 00 240 15 00 240 17 =(00 240 20 00 240 22 00 240 Injected 0.006 gram. 22 15 ? 22 30 i Dead. Experiment No. 34. Time: Pulsations REMARKS. min. sec. per minute. Normal ache 220 Pneumogastric nerves and cord cut. Injected intravenously 10 215 0.003 gram dried Cobra venom dissolved in 1 ¢. ec. distilled 30 215 water. 00) 210 3 00 215 500 215 8 00 225 11 00 225 14 00 225 I) Oo) 225 Killed by pithing. THE ACTION OF VENOMS UPON THE PULSE-RATH. 69 Summary and Conclusions of the Actions of Venoms on the Pulse-rate.—The results of this series of experiments indicate that the primary tendency of venoms is to cause an increase of the pulse-rate, that this tendency is greater after section of the pneumogastric nerves, and that it rarely occurs after conjoined section of the pneumogastric nerves and the upper or middle cervical region of the spinal cord. From the increased tendency to acceleration of the pulse-rate in poisoning by venom after section of the pneumogastric nerves we infer that there is some direct or indirect effect of the venom upon the pneumogastric centres by which an inhibi- tory influence is exerted, and which tends to neutralize the action bringing about acceleration. Since hastening of the pulse is a rare occurrence after conjoint section of the pneumogastric nerves and the cervical spinal cord, we think that the increase is due for the most part to some effect upon the accelerator centres in the medulla, whereby impulses are sent through (chiefly at least) those of the accelerator fibres which pass by the cord. ‘The increase of the pulse-rate which may occur after division of the nerves distributed to the heart, by section of the pueumogastric nerves and cervical spinal cord, must be dependent upon a direct action of the venom upon the heart muscle or its contained ganglia. The diminution in the heart beats must be due to a direct cardiac action, since it occurs after isolation of the heart, as above, from any central nervous influence. In these as in all other experiments which involve intravenous use of venoms we are liable to disturbing elements which do not trouble our explanations in dealing with other poisons. At any moment, anywhere in nerve-tissue or muscles, we may have abrupt and quite countless hemorrhages. How these may introduce con- flicting symptoms and modify results has already been pointed out by one of us many years ago." They make absolute constancy of effects quite improbable. Section I].—Tur Actions oF GLOBULINS ON THE PULSE-RATE. The Actions of the Venom Globulins on the Pulse-rate.—The actions of the venom globulins upon the pulse-rate appear to differ somewhat in quality from what is found in poisoning with pure venoms; there is a greater tendency to the primary increase in the pulse than with pure venoms, while the action by which this is brought about seems to differ. Of eleven experiments in which the amounts used represented the proportion of the respective globulins contained in the usual doses of venom given, six were made with the water-venom-globulin, two with the copper-venom-globulin, and three with the dialysis-venom-globulin ; all of these poisons, excepting in one experiment with the water-venom-globulin of the Ancistrodon, were derived from the venom of the Crotalus adamanteus. The water-venom-globulin seems to be the most active, and the copper-venom- * Researches on the Venom of the Rattlesnake. S. Weir Mitchell, 1861. 70 THE VENOMS OF CERTAIN THANATOPHIDESA. globulin the least so. Of the six experiments with the former, in four there was a primary increase in the pulse-rate followed by diminution, and in one case by a subsequent increase; in the other two there was a diminution from the first, the pulse regaining its normal frequency, or, as in one instance, rising above it. In both the experiments with copper-venom-globulin there was a primary increase followed by a diminution in one case, and in the other by a return of the rate to about the normal. In the three experiments with dialysis-venom-globulin, a primary increase occur- red. In two this was followed by a drop below normal, while in the other the rate remained above the normal. Experiment No. 35. Time: Pulsations REMARKS. min. sec. per minute. Normal a Bie 290 Injected intravenously 0.0012 gram water-venom-globulin 10 305 (= 0.015 gram dried venom) from the venom of the Cro- 20 310 talus adamanteus. 40 315 1 00 290 iL 3 270 3 00 270 5 00 210 at 00 270 ) OO 280 12 00 290 15; @) 300 13) CO — Bl 25 00 320 35 600 330 45 00 330 55 00 330 Killed. Experiment No. 36. Time: Pulsations REMARKS. min. sec. per minute. 5 Normal 5 ce 310 Injected intravenously the water-venom-globulin from 0.03 10 310 gram dried venom of the Crotalus adamanteus. 20 275 40 265 1 00 260 1 20 260 1 40 280 3° 40 290 Clot in canula. 7 40 310 9 40 310 Injected water-venom-globulin from 0.015 gram dried venom 10. 00 315 10 20 300 10 40 300 14 900 300 17 +00 300 20 00 300 30 00 300 Killed by pithing. THE ACTION OF VENOMS UPON THE PULSE-RATE. Ail Experiment No. 37. Normal Experiment No. 38. Normal Experiment No. 39. Normal Time: min. ll ] sec. 10 20 30 50 10 10 30 39 40 50 Time: min. sec. 10 20 30 50 1 00 1 30 3 30 5 6930 i 80 9 30 12 30 14 30 16 50 lt 30 19 30 21 30 26 00 28 00 30 00 BO) 115) 35 «(00 37 «(00 39 00 Time: min. sec. 30 50 0.0) 1 30 I 50 2 00 2 40 3 10 Pulsations per minute. 320 300 330 305 300 300 300 310 310 310 310 Pulsations per minute. 2710 290 295 295 260 255 260 265 260 265 265 260 260 270 275. 275 260 260 260 260 260 260 260 255 Pulsations per minute, 280 270 230 220? 280 260 180 260 280 REMARKS. Injected intravenously 0.0033 gram water-venom-globulin from the dried venom of the Crolalus adamanteus dissolved by the addition of a trace of sodic carbonate. Injected double dose. Injected double dose. Killed by pithing. REMARKS. Injected intravenously the water-venom-globulin from one minim of fresh venom of the Crotalus adamanteus. Clot in canula. Clot in canula. Animal killed by pithing. REMARKS. Injected intravenously the wafer-venom-globulin from 0.004 gram dried venom of the Ancistrodon piscivorus dissolved in 1 ec. ec. distilled water by the addition of a few crystals of sodie chloride. Injected a similar dose. 72 THE VENOMS OF CHRTAIN THANATOPHIDEA: Experiment No. 40. Time: Pulsations ‘ REMARKS. min. sec. per minute. Normal anes 312 Injected intravenously the water-venom-globulin from 0.015 0 46 gram dried venom of the Crotalus adamanteus. 10 314 20 33.0 30 360 1 30 316 2 30 304 5 30 324 10 30 354 14 30 360 Heematuria. 19 30 396 24 30 384 29°30 372 34 30 3712 42 30 372 47 30 372 52 30 372 57 30 360 67 30 316 T7 30 316 80 30 120 She sere Dead; ecchymoses in intestines ; blood fluid. Experiment No. 41. Time: Pulsations . REMARKS. min. sec. per minute. Normal ee 280 Injected intravenously 0.0012 gram copper-venom-globulin 10 285 from the dried venom of the Crotalus adamanteus: 20 290 30 285 50 280 2 50 270 4 50 270 6 50 260 8 50 260 10 50 255 iil §@ 260 1 620 280 18 20 280 Injected a similar dose. 18 30 300 18 40 290 18 50 285 19 00 280 20 00 285 22 +00 285 27 00 290 THE ACTION OF VENOMS UPON THE PULSE-RATHE. 13) Experiment No. 42. Time: Pulsations REMARKS. *min. sec. per minute. Normal suet 290 Injected intravenously 0.00225 gram copper-venom-globulin 10 290 from the dried venom of the Crotalus adamanteus. 30 305 00) 310 3 00 310 5 00 310 0.0 310 8 00 310 Clot in canula. 10 00 310 12 00 312 22 00 280 94 600 280 26 00 280 Injected 0.0045 gram. 26 10 285 26 20 290 26 30 290 27 «400 290 27 00 290 29° 00 280 31 00 270 34 00 250 39 00 295 41 00 290 43 00 285 45 00 285 52 00 295 - fxs} (OX) 295 Experiment No. 43. Time: Pulsations REMARKS. min. sec. _ per minute. Normal eens 290 Injected intravenously 0.0017 gram dialysis-venom-globulin 20 305 from the dried venom of the Crotalus adamanteus dissolved 40 305 in 1c. e. distilled water with a trace of sodic carbonate. 50 305 1 20 295 3 20 275 5 20 275 Animal broke loose. 18 20 280 18 23 oe: Injected 0.0034 gram dialysis-venom-globulin. 18 30 290 18 45 280 19 05 Bia Struggles. 1G) 5) 270 19) 55) 270 20 «25 270 Pile 138 22 00 : 30 00 oss Dead. 10 May, 1886. 74 THE VENOMS OF CERTAIN THANATOPHIDEA. Experiment No. 44. Time: Pulsations / REMARKS. min. sec. per minute. Normal Ebe 265 Injected intravenously dialysis-venom-globulin from the dried 20 280 venom of the Crotalus adamanteus. 30 290 1 00 270 1 20 270 1 40 270 2 00 265 2 40 280 3 40 280 5 40 285 6 20 285 6 50 285 7 20 300 T 50 300 8 50 300 9 20 300 ) oO 300 10 50 300 il 6&0 * 300 12 50 300 14 20 300 Experiment No. 45. Time: Pulsations REMARKS. min. sec. per minute. Normal Pica 270 Injected intravenously 0.0017 gram dialysis-venom-globulin 10 280 from the dried venom of the Crolalus adamanteus. 20 300 30 295 1 00 280 3 00 276 5 00 260 : e OO 250 Clot formed in canula. 16 00 Bak Injected 0.0034 gram. ly 30) 255 i as ae 17 40 275 18 30 260 20 30 270 Struggles. 23 3 260 28 30 260 43 30 270 Dane) 290 The Actions of the Venom Globulins on Animals with Cut Pneumogastrie Nerves.—In five experiments on animals with cut pneumogastric nerves—one with the water-venom-globulin, two with copper-venom-globulin, one with dialysis-cenom- globulin (all from the Crotalus adamanteus), and one with the water-venom-globulin of the Cobra—there was a tendency to a lowered pulse-rate, although in one experiment there was a primary increase, and in another a slight increase above the healthy number after repeated injections. The effects were generally less than in normal animal. THE ACTION OF VENOMS UPON THE PULSE-RA'TEH. 715 Experiment No. 46. Normal Experiment No. 47. Normal Experiment No, 48. Normal Experiment No. 49. Normal Time: min. sec. 10 20 1 00 1 40 3 40 5 40 Time: min. sec. 0 15 25 45 1 15 2 00 4 00 3 00 13 00 18 00 23 00 28 Time: min. sec. 20 40 i 1@ 3 I@) 5 10 7 O %) it) 23 10 23 40 24 15 Time: min. — CO D> > bo sec. 40 50 50 50 50 50 50 Pulsations REMARKS. per minute. 205 Pneumogastrie nerves cut. Injected intravenously 0.0011 220 gram water-venom-globulin from the dried venom of the 230 Crotalus adamanteus. 210 190 170 180 Clot in canula. Pulsations REMARKS. per minute. 324 Pneumogastric nerves cut. Injected intravenously water- venom-globulin from 0.035 gram dried Cobra venom dis- solved in 1 ¢. ec. distilled water. 312 318 264 300 304 276 288 310 319 Animal broke loose from canula. Pulsations REMARKS. per minute. 305 Pneumogastrie nerves cut. Injected intravenously 0.0012 300 gram copper-venom-globulin from the dried venom of the 288 Crotalus adamanteus. 285 285 285 300 290 310 Injected 0.0024 gram, 310 us a 310 Killed. Pulsations REMARKS. per minute. 300 Pneumogastric nerves cut. Injected intravenously 0.0012 300 gram copper-venom-globulin from the dried venom of the 300 Crotalus adamanteus. 300 300 300 300 300 300 16 THE VENOMS OF CERTAIN THANATOPHIDESA. Time: min, sec, 13° 50 Wey 5x) 16 10 16 20 16 30 16 45 17 45 19 45 21 45 23 45 25 45 26 45 27 00 Experiment No. 50. Time: min. sec. Normal ; 10 20 30 50 i) SO 4 20 G20 8 20 LO 20 12 20 17 50 18 20 18 40 19 00 1G) ils 19) 20 Bil Bo) 23 00 25 00 27 6000 29 00 34 00 34 630 38 30 41 00 47 00 49 00 Pulsations per minute. 300 300 300 300 300 300 300 300 300 270 270 270 2710 Pulsations per minute. 310 305 300 300 310 300 310 300 295 295: 300 300 300 310 320 530 320 310 310 310 310 305 300 290 , REMARKS. Injected 0.0024 gram. “ & “ Struggles. Animal killed by pithing. REMARKS. Pneumogastric nerves cut. Injected intravenously 0.0017 gram dialysis-venom-globulin from the dried venom of the Crotalus adamanteus. Struggles. Struggles. Injected 0.0034 gram. GG (a3 “cc Struggles. (7G “c Dead. The Actions of Venom Globulins on the Pulse-rate in Animals with the Pneumo- gastric Nerves and Cervical Spinal Cord OCut.—In four experiments in which the pneumogastric nerves and spinal cord in the middle cervical region were cut—one was made with the water-venom-globulin, one with the copper-venom-globulin, and two with dialysis-venom-globulin of the Crotalus adamanteus: in one experiment THE ACTION OF VENOMS UPON THE PULSE-RATE. TH there was a fall followed by a rise to the normal, and succeeded by a slight fall; in a second the pulse-rate always remained below normal, while in the third there was an almost inappreciable rise, this followed by a fall, and by an increase due to a further injection of the poison. ‘The last showed a slight fall, then a return to the normal. Experiment No. 51. Time : Pulsations REMARKS. min. sec. per minute. Normal ee 250 Pneumogastric nerves and cord cut. Injected intravenously 10 250 0.0011 gram water-venom-globulin from the dried venom 30 215 of the Crolalus adamanteus. 1 00 240 1 10 240 2 10 245 3 10 245 5 10 245 2 7 10 245 8) UM) 245 iit 1@ 250 15 10 250 17 40 245 19 00 240 21 00 240 23 00 240 .25 00 240 27 00 240) Killed. Experiment No, 52. Time: Pulsations REMARKS. min. sec. per minute. : Normal ba 0 255 Pneumogastric nerves and cord cut. Injected intravenously a 10 255 0.0048 gram copper-venom-globulin from the dried venom 20 240 of the Crotalus adamanteus. 40 250 1 00 250 ‘ 3 30 240 5 30 225 8 BO) 225 iit Si) 210 13 30 210 16 30 210 0) 204 Injected 0.0048 gram. ly 30) 210 18 00 210 20 00 210 24 00 195 26 00 180 28 00 180 830 00 180 32 00 180 34 00 187 Killed. 718 THE VENOMS OF CERTAIN THANATOPHIDEA. Experiment No, 53. Time: Pulsatious , REMARKS. min. sec. per minute. Normal BN ne 240 Pneumogastric nerves and cord cut. Injected intravenously 10 240 0.0017 gram dialysis-venom-globulin from the dried venom 30 245 of the Crotalus adamanteus. 1 00 230 3 00 220 6 00 220 8 30 220 10 30 230 Injected 0.0034 gram. 12 30 230 sf “ “ 12 50 220 13 10 225 13 30 210 13 50 210 1500 200 16 00 190 18 00 Some Dead. Experiment No. 54. Time: Pulsations REMARKS. min. sec. per minute. Normal Boe 300 Pneumogastrie nerves and cord cut. Injected intravenously 10 290 0.0068 gram dialysis-venom-globulin from the dried venom 20 290 of the Crotalus adamanteus. 30 300 40 300 Tremors. 50 300 1 00 300 1 IO 300 1 20 300 Clot formed in canula. 9 00 es Dead. A review of the results of these experiments with the globulins on the pulse-rate in normal animals indicates that water-venom-globulin is the most potent, and the copper-venom-globulin the least so. With the former there occurred in four of the six experiments a primary increase followed by a fall, while in the other two there was a diminution from the first. In experiments with the copper-venom-globulin and dialysis-venom-globulin there was always a primary increase, and in four out of the five experiments this was followed by a decline. After section of the pneumogastric nerves a primary increase (due probably to some accidental cause) occurred in one out of the five experiments, in two of the other four there at first was no appreciable change, and then a diminu- tion, while in the remaining two there was a lessening of the rate from the time of injection. These results suggest that the increase of the pulse-rate, which occurred in animals with intact vagi, was in some degree at least dependent upon an influence exerted through the pneumogastric centres and nerves. It will be observed that we here have results which are directly opposed to what we have seen with pure venom; that is a lessened tendency to the primary increase of the 2 THE ACTION OF VENOMS UPON THE PULSE-RATE. 79 pulse after section of the pneumogastric nerves. If the increase in the pulse-rate in normal animals is due for the most part to excitation of the accelerator centres, whereby impulses are generated which pass chiefly through the accelerator fibres running in the spinal cord, it would seem probable that the accelerator impulses induced by the globulins take for the most part the course of the fibres through the pneumogastric nerves, but are much feebler than the impulses which are gene- rated by the pure venoms, and which take their path chiefly through the fibres in the spinal cord. After section of both the pneumogastric nerves and cervical spinal cord, we found in all of our experiments a diminution in the heart-beats; this must be due to a direct action of the globulin upon the heart. It therefore seems probable that the globulins cause a primary increase of the pulse by an excitation of the accelerator centres, whereby impulses are conveyed principally by the accelerator fibres passing through the pneumogastric nerves; and a diminution of the heart beats by a direct action on the heart. Section JI1].—Tur Actions or VENOM PEPTONES UPON THE PULSE-RATE The Action of Venom Peptones on the Pulse-rate-—In seven experiments made with peptone on normal animals—four with the peptone from the venom of the Crotalus adamanteus ; one with that of the Ancistrodon piscivorus ; and two with that of the Cobra—we find results which vary and which resemble closely those obtained by the administration of pure venom. In three experiments there was a primary increase of pulse followed generally by a diminution; in three the pulse remained below normal; while with Ancistvodon peptone there was a primary fall of rate followed by a rise. The differences in the results, as in previous experiments, do not seem to depend at all upon the dose or the variety of venom from which the peptone was obtained. Experiment No, 55. Time: Pulsations REMARKS min. sec. per minute. Normal hone 280 Injected intravenously the peptone from 0.015 gram dried 20 102 venom of the Crotalus adamanteus. 30 190 40 190 50 190 1 00 180 3 00 190 6 00 340 Struggles. 11 090 285 Struggles. Broke loose. 49 00 oe Dead. 80 THE VENOMS OF CERTAIN THANATOPHIDE A. Experiment No, 56. Time : Pulsations / REMARKS. min. sec. per minute. Normal re 225 Injected intravenously the peptone from 0.03 gram dried 10 260 venom of the Crotalus adamanteus. 30 260 1 00 285 2 00 270 5 00 260 900 250 Killed by pithing. Experiment No. 51. Time: Pulsations REMARKS, min. sec. per minute. — Normal eee 280 Injected intravenously the peptone from 0.015 gram dried 30 280 venom of the Crotalus adamanieus. 1 00 270 1 30 285 4 30 270 10 30 270 Injected double the amount. 10 50 270 > mt id 270 11 30 270 Experiment No. 58. Time: Pulsations REMARKS. min. sec. per minute. : , Normal Seas 270 Injected intravenously the peptone from 0.015 gram dried 10 270 venom of the Crotalus adamanteus. 20 2710 30 270 1 00 280 1 30 290 2 00 290 2 30 290 3 00 260 5 00 260 Injected a similar quantity. 5 20 260 5 40 260 6 00 260 6 30 255 7 00 250 Injected double the quantity. 7 30 240 8 00 260 Experiment No. 59. Time: Pulsations REMARKS. min. sec. per minute. Normal SUES 300 Injected intravenously the peptone from 4.05 gram dried 10 175 venom of the Ancistrodon piscivorus. 30 110 10.0 285 eA) 290 1 40 300 2 00 310 THE ACTION OF VENOMS UPON THE PULSE-RATHE. 81 Experiment No. 60. Normal Experiment No. 61. Normal 11 Time: min. 2 C2 SS oS MN Or pO —_ Sec. 20 40 40 50 Time: min. sec. Co 6 11 16 16 17 18 18 9) 19 19 19 20 20 10 20 30 40 00 20 20 20 20 20 50 20 50 00 10 30 50 10 30 Time: min. sec. 3 8 10 10 10 11 11 12 12 12 May, 1886, 10 15 30 50 20 20 20 40 50 20 40 00 20 40 Pulsations per minute, 340 315 340 348 340 340 340 Pulsations per minute. Pulsations per minute. 290 300 310 240 245 225 97 72 300 300 105 85 120 90 REMARKS. Injected peptone from 0.01 gram venom, Killed. REMARKS, Injected intravenously the peptone from 0.005 gram dried Cobra venom. Struggles, Convulsive twitchings. Blood is asphyxiated ; no respiration Killed, REMARKS. Injected intravenously the peptone from 0.06 gram dried Cobra venom. Tonic convulsions. Convulsive twitchings; asphyxiated blood; respiration ceased. Dead. 52 THE VENOMS OF CERTAIN THANATOPHIDESR. The Actions of Venom Peptones on Animals in which the Pnewmogastric Nerves had been Cut.—¥our experiments were made with the peptone on animals the pheumogastric nerves of which had been previously cut. In three of the four there was a primary increase in the pulse, while in the fourth there was a temporary diminution followed by a rise above normal. These experiments are in accord with those made with pure venom, and indicate a greater tendency to primary pulse frequency after section of the pneumogastric nerves. One of the above experiments was made with the peptone from the Crotalus adamanteus ; one with the Ancistrodon piscivorus ; and two with the Cobra. Experiment No. 62. Time: Pulsations REMARKS. min. sec. per minute. Normal 9 285 Pneumogastric nerves cut. Injected intravenously the peptone 10 285 from 0.015 gram dried venom of the Crotalus adamanteus. 30 285 40 300 Struggles. 50 300 1 00 300 1 20 300 I aX) 300 8 8 . sil 8 30 330 Struggles. 15 30 345 20 30 325 25 30 315 30 30 315 35 00 315 40 00 315 45 00 270 50 00 255 62 00 285 Killed. Experiment No. 63. Time: Pulsations REMARKS, min. sec. per minute. Normal ee 295 Pneumogastric nerves cut. Injected intravenously the peptone 30 315 from 0.015 gram dried venom of the Ancistrodon pisci- 40 315 vorUs. 50 310 1 00 310 1 20 315 1 40 320 2 00 330 2 30 310 2 40 102 2 50 150 3 00 240 4 20 000 Dead. THE ACTION OF VENOMS UPON THE PULSE-RATE. 83 Experiment No. 64. Time: Pulsations REMARKS, min. sec. per minute, Normal oat 230 Pneumogastric nerves cut. Injected intravenously the peptone 10 235 from 0.005 gram dried Cobra venom. 20 240 40 230 1 00 230 1 30 230 3 30 230 5 30 230 7 30 230 9 30 220 10 00 220 21 30 225 Twitchings. 25 30 230 26 30 235 Killed. Experiment No. 65. Time: Pulsations REMARKS. min. sec. per minute. Normal 16 ¢ 265 Pneumogastric nerves cut. Injected intravenously the peptone 10 265 from 0.006 gram dried Cobra venom. 20 255 40 260 1 00 260 3 00 270 5 00 270 16 00 275 34 00 Biota Dead. The Actions of Venom Peptones upon the Pulse-rate of Animals after Section of the Pnewmogastric Nerves and Cervical Spinal Cord.—Six experiments made on animals in which the heart was cut off from central nervous influence by section of the pneumogastric nerves and section of the spinal cord in the middle cervical region gave uniform results. Three were made with peptone from the venom of the Crotalus adamanteus ; two with the peptone from the Ancistrodon piscivorus, and one with that of the Cobra. In all of these experiments there was a diminu- tion of the pulse-rate, and usually this was well marked. ‘These results are also in accord with what was found in experiments with pure venom. Experiment No. 66. Time: Pulsations REMARKS. min. sec. per minute. Normal Ss 3H 0 200 Pneumogastric nerves and cord cut. Injected intravenously 10 185 the peptone from 0.015 gram dried venom of the Crotalus 20 195 adamanteus. 40) 195 1 00 190 3 00 160 13 00 195 15 00 200 84 THE VENOMS OF CERTAIN THANATOPHIDEA, Time: Pulsations REMARKS min. sec. per minute, , 17 00 190 Injected peptone from 0.03 gram dried venom. 17 30 Sees 19 30 187 21 30 190 23 30 180 25 ©30 170 31 630 165 34 30 BIEnG Dead. Experiment No. 67. Time: Pulsations : REMARKS. min. sec. per minute. Normal Apes 282 Pneumogastric nerves and cord cut. Injected intravenously 0 aad the peptone from 0.015 gram dried venom of the Crotalus 15 276 adamanteus. 20 ies 30 264 1 00 264 1 30 246 2 00 234 Experiment No. 68. Time: Pulsations REMARKS, min. sec. per minute. Normal aid 6 324 Pneumogastric nerves and cord eut. Injected intravenously 0 soi the pepione from 0.015 gram dried Cobra venom. Ii 300 40 318 1 40 276 5 40 272 10 40 276 15 40 306 Me OD } Injected a similar dose. 16 06 tases 1 15) 306 1G 2) 300 0,0 276 18 00 270 23 00 oe Dead. In the above series of experiments with venom peptones we find results which agree with those in which the pure venoms were used. We conclude, therefore, that the peptones cause a primary increase and a secondary diminution of the pulse-rate, and that they occasion primary hastening of the heart beat by excita- tion of the accelerator centres in the medulla, and that the impulses are carried through fibres passing chiefly by the spinal cord. ‘This increase is more marked after section of the pneumogastric nerves; thus suggesting that this principle has some direct or indirect effect upon the pneumogastric centres, tending to slow the action of the heart and to neutralize the accelerator influence. Peptones cause the diminution of the heart beat by a direct action on that organ. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 85 CHAPTER VIII. THE ACTION OF VENOMS AND THEIR ISOLATED GLOBULINS AND PEPTONES UPON THE ARTERIAL PRESSURE. Section I.—Purr VENOM. THE experiments made on the blood pressure with venoms and their isolated poisons were all made on rabbits. ‘The manometer tube was connected with one of the carotid arteries, and the injections were always made into the external jugular vein unless otherwise noted. Eighteen experiments were performed with the venoms of different species of serpents, and in all of them there was a distinct lowering of the blood-pressure. It fell immediately after the injection, and indeed sometimes before injection was complete, and the fall was generally so marked as to indicate a most profound action of the poison upon some part or parts of the circulatory apparatus. If the dose be not immediately fatal the pressure gradually rises, but finally undergoes a more or less steady decline to death. At other times the pressure sinks without subsequent rise until death ensues. The tendency in Cobra poisoning is to a decided rise of pressure following the primary fall. In five out of six experiments with this venom the primary fall was followed by a rise which went above the normal. Of the eighteen experiments, five were made with the venom of Crotalus ada- manteus, in two of which the poison was given hypodermatically; two with that of the Crotalus horridus ; two with the venom of Ancistrodon piscivorus ; one each with the poisons Ancistrodon contortrix, Crotalophorus miliarius, and Daboia Russellic ; and six with the venom of the Cobra. In all cases ether was given freely to the animal poisoned. Action of the Pure Venoms upon the Arterial Pressure in Normal Animals. - Experiment No. 1. Time: Pressure REMARKS. min. sec. m. Mm. Normal Beanie 126 Injected into the thigh of a large rabbit 1 drop of fresh venom 20 126 from the Crotalus adamanteus. 40) 126 00 124 120 192 1 40 120 2 00 118 Clot formed in canula. 5 00 114 86 THE VENOMS OF CERTAIN THANATOPHIDE SA. Time: min. 7 8 9 10 11 12 13 20 21 23 25 26 Experiment No. 2. sec. 00 00 00 00 00 00 00 00 30 00 00 Time: min. Normal be = Experiment No. 3. sec. 20 40 00 20 40 00 Time: min. sec. Normal bp po ke bo Experiment No. 4. 10 20 30 40 50 00 20 40 00 20 40 Time: min. sec. Normal Doe ee 5 10 20 30 40 00 20 40 10 Pressure m. m. 84 84 84 82 82 78 82 52 56 64 56 Pressure m. m. 144 144 138 136 130 130 126 Pressure m. m. 70 58 54 60 68 68 68 68 62 58 54 44 Pressure m. mM. 110 76 72 64 62 60 58 58 56 54 REMARKS. Struggles followed by death. REMARKS. Injected into the thigh of a rabbit 3 drops of fresh venom from the Crotalus adamanteus. Struggles. Animal tore loose from the canula. REMARKS. Injected intravenously 0.003 gram dried venom of the Cro- talus adamanieus in 1 e. c. distilled water. Death. REMARKS. Injected intravenously 0.003 gram dried venom of the Cro- talus adamanteus dissolved in 1 c¢. ec. distilled water. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 87 Time: min. sec. 4 00 7 00 8 00 8 10 8 20 8 30 8 40 850 10 30 13 00 13 30 13 50 14 00 16 00° 16 05 16 30 17 (00 Experiment No. 5. Time: min. Normal (Cos ll Experiment No. 6. sec. 10 20 30 40 00 20 40 55 00 00 00 Time: min. Normal oo Tow Be ee _ sec. 10 20 30 40 50 00 10 20 30 40 40 40 40 40 10 Pressure m. m. 45 40 110 94 74 78 76 60 56 64 50 50 48 44 Pressure m. m. 124 100 60 96 34 70 56 48 44 38 32 Pressure m. m. 104 84 68 74 80 18 70 64 60 56 52 48 40 44 46 42 38 REMARKS. Struggles. Injection as before. Dead. Heart in complete diastole. Hechymoses in pericar- dium and peritoneum. Blood incoagulable. REMARKS. Injected intravenously 0.015 gram dried venom of the Cro- talus adamanteus dissolved in 1 ¢. c. distilled water. Struggles. Pulse feeble. Dead. REMARKS. Injected intravenously 0.015 gram dried venom of the Cro- talus horridus dissolved in 1 ec. c. distilled water Convulsions. Dead. Some ecchymoses; blood fluid. 88 Experiment No. 7. Normal Experiment No, 8. Normal Experiment No. 9. Normal THE VENOMS OF CERTAIN THANATOPHIDESA. Time: min. sec. Oo OO CO ST co OF WH a S 5 10 20 30 40 00 00 30 30 00 30 00 00 Time: min. sec. BS Be) We) RS) Ie JS tS 20 30 40 00 30 50 10 30 00 00 Time: min. sec. TO SS SD Or or Or Or GD bb Fe CFS 10 20 30 00 30 00 30 00 00 3 39 45 05 15 20 45 05 Pressure ’ m. m. 110 76 76 . U8 70 60 44 42 40 36 32 26 20 10 Pressure m. m. 138 80 64 74 84 76 60 74 74 84 Pressure m. m. 134 108 72 70 70 12 74 70 70 70 68 16 70 60 62 66 60 52 REMARKS. Injected intravenously 0.015 gram dried venom of the Cro- talus horridus dissolved in 1 ¢. e. distilled water. Animal broke loose from mouth-piece, and was firmly held and refixed. Dead. Respiration failed before the heart. REMARKS. Injected intravenously 0.004 gram dried venom of the Ancis- trodon piscivorus dissolved in 1 ec. ce. distilled water. Convulsions. Injected as above. Struggles. Killed by pithing. REMARKS. Injected intravenously 0.004 gram dried venom of the Ancis- trodon piscivorus dissolved in 0.5 c. c. distilled water. Injection repeated as before Convulsive movements. THE ACTION Time: min. sec. eels 7 20 7 3 7 45 qT 55 8 05 8 15 8 25 OF VENOMS UPON ARTERIAL PRESSURE. 89 Pressure m. m. 54 72 70 66 104 120 100 86 Experiment No. 10. Time: Normal min. Soot OOOO fF DW = — sec. 10 30 00 30 00 2) 30 30 50 30 50 50 00 Pressure m. m. 96 70 76 72 72 72 70 48 48 42 50 50 46 Experiment No. 11. Normal Time: min. co —T Ort bw bw bt ee sec. bo bt) bo wo Soa] So Pressure m. m. 170 122 120 136 116 150 84 108 106 80 84 70 74 80 Experiment No. 12. Time: Normal 12 min. June, 1886. sec. 10 20 30 40 Pressure m. m. 130 110 102 96 88 REMARKS. Animal died in a few minutes. REMARKS. Injected intravenously 0.003 gram dried venom of the Ancis- trodon contortrix dissolved in 1 c. ¢. distilled water. Injection repeated. Struggles. Injection repeated. Dead. Heart flabby~ blood incoagulable; no ecchymoses. REMARKS. Injected intravenously 0.003 gram dried venom of the Crota- lophorus miliarius dissolved in 1 ec. ¢. distilled water. Injection repeated as before. . Killed by pithing. REMARKS. Injected intravenously 0.003 gram dried Cobra venom dis- solved in 1 ¢. ec. distilled water. 90 Experiment No. 13. Normal Experiment No. 14. Normal Experiment Normal THE VENOMS OF CERTAIN THANATOPHIDESA, Time: min. (JU OR Ol sec, 50 00 10 20 30 40 00 20 20 Time: min. oq = = sec. 10 30 00 20 20 20 Time: min. sec. 1 3 8 10 15 17 18 19 COF WNW SO Ee DH ho ee 00 00 00 00 00 00 00 00 No. 15. Time: min. sec. 20 40 00 30 00 00 00 00 00 00 Pressure m. m. 70 78 70 52 52 50 38 34 Pressure m. m. 140 150 140 134 134 140 148 Pressure m. m. 13 122 120 130 138 106 66 48 Pressure m. m. 145 142 135 128 130 140 150 143 138 158 135 REMARKS, Convulsions. Dead. Heart in systole; blood clotted; no ecchymoses. REMARKS. Injected intravenously 0.003 gram dried Cobra venom dis- solved in 1 c. c. distilled water. Death from hemorrhage ; artery torn. REMARKS. Injected intravenously 0.003 gram dried Cobra venom dis- solved in 1 c. ¢. distilled water with a few crystals of sodic chloride. Dead. REMARKS. Injected intravenously 0.003 gram dried venom of the Cobra dissolved in 1 ¢. c. distilled water. Respiration ceased; artificial respiration used. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 91 Experiment No. 16. Normal Experiment No. 17. Normal Experiment No. 18. Normal Time: min. sec. moO CO Te eS eS eR — 20 30 40 00 20 40 40 40 40 10 00 Time : min. sec. ho Ht ee DATADA Do — S LO 10 20 30 40 00 30 00 10 10 40 10 10 20 50 30 20 Time: min. Dee sec. 10 11 15 20 30 40 50 00 15 00 Pressure m. m. 120 108 96 88 90 82 96 94 100 122 156 Pressure m. m. 90 94 66 84 90 88 86 78 74 90 96 92 72 60 32 24 Pressure m. m. 110 112 80 64 46 30 o4 46 46 REMARKS. Injected intravenously 0.005 gram dried Cobra venom dis- solved in 1c. c. distilled water with a little sodic chloride and filtered. A clot was probably beginning to form in the canula, and no dependence is to be placed upon the after record. Struggles. Clot in canula. Dead. REMARKS. Injected intravenously 0.015 gram dried Cobra venom dis- solved in 1 c. c. distilled water. Dead. REMARKS. Injected intravenously 0.0045 gram dried venom of the Daboia Russelliz dissolved in 0.5 ec. e. distilled water. Pressure falling. Violent general convulsions. Dead. Heart in diastole; no ecchymoses; after twenty-four hours the blood is still fluid. Artificial respiration was used in this experiment from the beginning. 92 THE VENOMS OF CERTAIN THANATOPHIDESA. This single experiment confirms the statements of Fayrer and of Wall in regard to the convulsivant power of Daboia. The spasms are not due to defect of oxygen, as they arise early and occur despite the use of artificial respiration. _ Ancistrodon venom seems to have the same capacity to produce convulsions. The Action of Pure Venoms on the Blood Pressure of Animals with Cut Pnewmo- gastric Nerves.—After section of the pneumogastric nerves, including the depressor fibres, we find that the same alterations occur in the blood pressure as in normal animals. Nine experiments were made altogether: two with the venom of the Crotalus adamanteus ; one with the Crotalus horridus ; two with the Ancistrodon piscivorus ; one with the Ancistrodon contortria ; and three with the Cobra. Experiment No. 19. Time: Pressure REMARKS. min. sec. m. m. Normal flies 96 Injected intravenously 0.003 gram dried venom of the {'ro- 10 74 talus adamanteus dissolved in 1 e. c. distilled water. 30 68 O00 68 It s0 16 2 00 64 2° 30 60 5 RR Ad 7 00 44 8 30 48 Injected the same as above. 8 40 42 9 00 46 9 10 44 9 20 44 9 8 44 Dead. Experiment No. 20. Time: Pressure REMARKS. min. sec. m. m. Normal Ane 130 Injected intravenously 0.015 gram dried venom of the Cro- 10 100 talus adamanteus dissolved in 1 e. c. distilled water. 20 90 30 96 40 16 50 56 1 00 50 lO 38 1 20 32 1 30 22 Dead. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 93 Experiment No. 21. Time: min. Normal come by bp ee Experiment No. 22. sec. Time: min. sec. Normal 1 Experiment No. 23. Time: min. sec. Normal 10 20 30 40 50 1 00 1 20 S00) 5 50 6 00 8 00 10 00 12 00 14 00 19 00 LS 10 19 20 LO 30 19 40 — o 8 10 20 30 40 00 20 40 50 Pressure m. m. 144 146 124 124 94 80 70 56 54 54 44 12 Pressure m. m. 90 76 o4 44 50 54 44 24 Pressure m. m. 110 84 64 68 78 76 72 66 52 64 86 100 74 70 66 70 70 52 98 68 90 REMARKS. Injected intravenously 0.015 gram dried venom of the Cro- talus horridus dissolved in 1 ¢. ¢. distilled water. Struggles. Violent struggles. Dead. REMARKS. Injected intravenously 0.0035 gram dried venom of the Ancis- trodon piscivorus dissolved in 1 ¢. ¢. distilled water. Convulsions. Dead. REMARKS. Injected intravenously 0.003 gram dried venom of the Ancis- trodon piscivorus dissolved in 1 c. e. distilled water. Struggles. Struggles. Convulsions. Injection repeated, using the same amount. 94 THE VENOMS OF CERTAIN THANATOPHIDES, Time: Pressure REMARKS. min. sec. m.m. , 20 00 90 20 20 72 21 20 60 23 20 56 | 25 20 60 | 28 20 70 33 20 70 38 20 66 | 44 20 58 Third injection, same amount. 44 30 50 44 40 48 44 50 44 45 00 BG. 45 10 26 Dead. Experiment No, 24. Time: Pressure REMARKS min. sec. m. m. Normal ime 154 Injected intravenously 0.003 gram dried venom of the Ancis- 10 114 trodon contortrix dissolved in 1 ¢. c. distilled water. 20 86 30 76 40 82 00 84 15.0) 80 4 20 98 7 00 100 Injection repeated. t OS 138 Struggles. T 10 110 i 2 106 7 40 108 8 00 98 8 30 88 9 00 80 Il sO 78 Third injection. | 140 76 12 00 70 12 30 66 13 00 68 13 30 68 19 00 72 Fourth injection. 19 20 54 19 40 50 20 20 44 21 50 40 22 50 38 Dead. Heart in diastole; blood remains fluid; muscles all respond to electrical irritation; motor nerves react feebly. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 95 Experiment No. 25. Time: min. sec. Normal 1 1 3 6 10 14 16 Experiment No. 26. 10 30 00 ~30 30 30 30 30 30 Time: min. sec. Normal Experiment No. 27. Time: min. sec. Normal : 0 10 20 1 00 2 00 4 00 10 15 10 30 00 30 30 30 30 30 9 3) 30 Pressure m. Mm. 148 146 136 140 140 150 150 146 152 156 Pressure m. m. 134 136 126 118 118 132 136 136 136 136 188 Pressure m. m. 130 130 118 118 115 REMARKS, Injected intravenously 0.003 gram dried Cubra venom dis- solved in 1 ¢. e. distilled water with a few crystals of sodic chloride and filtered. Clot formed in canula. Animal killed. REMARKS. Injected intravenously 0.006 gram dried Cobra venom pre- pared as in the foregoing experiment. Convulsive movements; asphyxia; respiration ceased in three minutes, REMARKS. Tnjected intravenously 0.003 gram dried Cobra venom dis- solved in 1 c. c. distilled water. Clot in canula. Dead from asphyxia The Action of Pure Venoms on the Blood Pressure of Animals in which the Cervical Spinal Cord had been Divided.—Upon section of the spinal cord in the upper cervical region, by which the influence of the vaso-motor centres in the medulla is practically destroyed, the primary fall of pressure from venom is gener- ally very slight, and after this diminution there is a secondary rise which may go above the normal. was a rise of pressure for a moment at the time of injection; in one experiment with Crotalus horridus, in which a somewhat larger dose was used than in the others, there was a distinct rise of pressure a few seconds after injection, followed by a fall; and in the experiment with the Cobra the pressure never went below In one experiment with Crotalus adamanteus venom there 96 THE VENOMS OF CERTAIN THANATOPHIDEA. the normal, but in a few moments a rise occurred which continued to increase for half an hour, when the animal’ was killed. In this series we observed a marked difference from the preceding (unless the dose had been immediately toxic), since the profound primary fall of pressure was not observed, excepting in a very slight degree if at all; we found, however, that the ultimate fall of pressure still occurred, save in the case of the Cobra. Eight experiments were made: two with Crotalus adamanteus ; one with Cro- talus horridus ; three with Ancistrodon piscivorus ; one with Ancistrodon contortria, and one with Cobra venom. Experiment No. 28. Time: Pressure REMARKS. min. sec. m,. m. 5 Normal Bet 62 Injected intravenously 0.003 gram dried venom of the Cro- 9 70 talus adamanteus dissolved in 1 ¢. ec. distilled water. 20 56 30 56 40 58 1 00 58 1 20 56 i SO 48 1 40 46 2 00 44 2 20 40 2 40 38 5ee20 36 6 50 36 ioe) 36 8 00 ees Dead. Experiment No. 29. Time: Pressure REMARKS. min. sec. m. m. Normal 0 66 Injected intravenously 0.006 gram dried venom of the Cro- 5 60 talus adamanteus dissolved in 1 c. ec. distilled. water. 10 58 2 OOS 30 64 40 62 50 58 1 00 56 1 30 48 2 30 40 3 3 60 4 30 56 5 30 40 (oe) 00 Sine Dead. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 97 Experiment No. 30. Time: min. sec. Normal 5 20 40 x 00 20) 3 20 5 20 7 20 9 20 10 50 12 50 14 50 Gia ie (085) 17 35 18 00 Experiment No. 31. Time: min. sec. Normal : 10 20 30 40 50 00 00 00 00 00 oar we Experiment No. 32. Time: min. sec. Normal ‘ 10 20 30 40 50 Experiment No, 33. Time: min. sec. Normal , 10 20 30 40 56 3) 20 om 00 8 00 13. June, 1886, Pressure REMARKS, m. m. 30 Injected intravenously 0.015 gram dried venom of the Cro- 30 talus horridus dissolved in 1 ¢. ce. distilled water. 46 42 38 26 26 26 24 30 30 28 26 26 10 Conjunctival reflexes gone. Dead. Pressure REMARKS. m. m. 56 Injected intravenously 0.007 gram dried venom of the Ancis- 50 trodon piscivorus dissolved in 1 ec. c. distilled water. 46 46 40 34 30 22 28 26 18 Dead. The cord proved not to have been completely cut—a few fibres of the posterior columns remaining undivided. Pressure REMARKS, m. m. 58 Injected intravenously 0.003 gram dried venom of the Ancis- 54 trodon piscivorus dissolved in 1 e¢. ¢. distilled water. 40 32 Convulsions. 26 16 Dead. Pressure REMARKS. m. m. 46 Injected intravenously 0.003 gram dried venom of the Ancis- 38 trodon piscivorus dissolved in 1 ¢. e. distilled water. 40 38 40 48 38 30 28 Dead. 98 THE VENOMS OF CERTAIN THANATOPHIDEA. Experiment No, 34. Time: Pressure REMARKS. min. sec. mm. , Normal a6 9 52 Injected intravenously 0.003 gram dried venom of the Ancis- 10 44 trodon contortria dissolved in 1 ¢. e. distilled water. 20 46 40 50 1 00 54 3 OO 46 5 00 36 Tt OW 34 9 00 34 11 00 30 13 00 30 15 00 30 17 00 30 19 00 3 21 00 30 23 «00 30 25 00 32 27 00 32 29 00 34 56 00 34 61 00 32 75 00 30 Killed by pithing. Experiment No, 35. Time: Pressure REMARKS. min. sec. m. m. Normal Par 42 Injected intravenously 0.003 gram dried venom of the Cobra 10 46 dissolved in 1 ¢. c. distilled water with a few crystals of 30 44 sodie chloride and filtered. 1 @0 42 3 00 46 6 00 48 8 OW 46 12 00 50 15 30 50 18 3 52 21 30 _ 54 24 30 56 27 30 54 Injected the same as the foregoing. 27 40 56 28 00 64 28 30 68 30 30 78 Clot formed in canula. Killed animal by pithing. The Action of Pure Venoms on the Blood Pressure of Animals in which the Pneumogastric, Depressor, and Sympathetic Nerves and Spinal Cord have been Severed.—Since we found in the last series of experiments that after section of the cord there did not occur such a decided primary fall of pressure, it seemed obvious that the fall of pressure must be due, in major part at least, to a toxic depression THE ACTION OF VENOMS UPON ARTERIAL PRESSURE 99 of the vaso-motor centres. A fall of pressure does, however, ultimately occur, and, excepting in the case of the Cobra, increases until death ensues In seven other experiments, supplementary to the above, in which we made section of the pneumogastric, depressor, and sympathetic nerves in the neck, and section of the spinal cord in the middle or upper cervical region, thus cutting off both the heart and capillaries from centric nervous influence, we obtained results which were practically the same Three of. these experiments were made with the venom of the Crotalus adaman- teus, one with that of the Crotalus horridus ; one with the Ancistrodon piscivorus ; one with the Ancistrodon contortrax, and one with the Cobra. Experiment No 36. Time Pressure REMARKS, min. sec. m. m. Normal a6 62 Injected intravenously 0003 gram dried venom of the Cro- 10 56 talus adamanteus dissolved in 1 ¢ © distilled water. 20 46 30 56 40 52 1 00 46 120 40 1 40 36 2 00 30 2 20 24 Dead. Heart arrested in diastole; blood incoagulable; a few ecchymoses in peritoneum. The section of the cord was not quite complete. Experiment No. 37. < Time: Pressure REMARKS. min, sec. m m. Normal é 48 Injected intravenously 0.003 gram dried venom of the Cro- 10 46 talus adamanteus dissolved in 1 ¢. ¢. distilled water. 20 48 30 46 1 00 48 1 20 46 3 20 40) 3 50 32 250 36 7 50 30 Dead. Heart arrested in diastole; blood incoagulable; no ecchymoses in serous tissues Expervment No 38. Time: Pressure REMARKS. min. sec. m. m. Normal OSS 3 Injected intravenously 0.003 gram dried venom of the Gro- 5 32 talus adamanteus dissolved in 1 ¢. c. distilled water, 10 28 20 28 30 26 40 28 50 28 100 Experiment No. 39. Normal Experiment No. 40. Normal KHxperiment No. 41. Normal THE VENOMS OF CERTAIN THANATOPHIDESA, Time: min. sec. 1 00 IL aio) 2 00 atl) 9 30 Time: min. Co S\N ll 8 12 Time. min. sec. 20 30 40 1 00 i a 0) 4 20 *6 20 8 20 10 20 12 20 15) 2X0) 18 20 21 20 Time: min. sec. 10 20 40 sec. 10 20 30 40 00 20 40 00 00 00 00 00 Pressure REMARKS. m. m. at 27 27 22 , Dead. Heart arrested in diastole; blood incoagulable , ecchymoses well-marked in peritoneum and pericardium ; intestines congested; 50 c.c serum in peritoneal cavity. Section of cord complete, except anterior columns. Pressure REMARKS. m,. m. tt Injected intravenously 0.015 gram dried venom of the Cro- 44 talus horridus dissolved in 1 c. c. distilled water. 56 62 60 52 44 44 38 30 99 Dead. Pressure REMARKS. m. m,. 46 Injected intravenously 0.003 gram dried venom of the Ancis- 40 trodon piscivorus dissolved in 1 ¢. c. distilled water. 48 Muscular movements. 44 44 40) bo bw bw oo a oo Dead. Blood is incoagulable; no ecchymoses in serous membranes. : Pressure REMARKS. m. m. 64 Injected intravenously 0.003 gram dried venom of the Ancis- 56 trodon contortria dissolved in 1 ¢. c. distilled water. 54 48 THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 101 Time: Pressure REMARKS. min. sec. m. m. 1 00 42 1 30 40 2 00 42 4 00 50 7 00 56 900 54 1l 00 48 13 00 48 15 00 48 17 +00 48 20 00 48 22° 00 48 Injected 0.006 gram. 22 15 38 22. 30 32 Dead. Experiment No, 42. Time: Pressure REMARKS. min. sec m. m. Normal ne ee 56 Injected intravenously 0.003 gram dried venom of the Cobra 10 60 dissolved in 1 c. c. distilled water and a few crystals of 30 56 sodie chloride and filtered. 1 00 52 3 00 48 5 00 48 8 00 52 11 00 58 14 00 60 19 00 62 Animal killed by pithing. To recapitulate the actions of pure venoms upon the arterial pressure—we find that the injection of venom subcutaneously causes a progressive fall of blood pressure; when injected intravenously, there is a sudden and decided fall of pressure, which may be immediately followed by death, or by a gradual rise, to be in turn succeeded by a decline with feeble pulse as death approaches. In the Cobra there is a tendency to a rise of pressure, which may go above the normal as death appears. After section of the pneumogastric nerves and its depressor fibres we find no alterations in the results obtained in normal animals, but when section of the cord is made in the middle or upper cervical region by which the vaso-motor centres in the medulla oblongata are practically destroyed, or when accompanying this section the nerves in the neck and the spinal cord in the middle cervical region are also cut, thus practically isolating the vaso-motor centres in the medulla and cutting off all central nervous connection with the heart, we find that the primary profound diminution of pressure is not so marked. ‘There may even appear to be a slight tendency on the part of the arterial pressures to rise above the normal just before death. Even after section of the spinal cord, as above, we find in Cobra the increase of pressure occurring before death as in normal animals. These results indicate that the primary positive failure of pressure is due chiefly 102 THE VENOMS OF CERTAIN THANATOPHIDESA. to a depressant action of the venom upon the vaso-motor centres in the medulla oblongata, and slightly upon the heart. ‘The tendency to a rise of pressure, as well as the ultimate fall, must be due to some action upon the heart itself or the general systemic capillaries. It seems probable that the rise of pressure in these experi- ments is of capillary origin since the pulse-curves do not indicate increased heart power, and we have already had reason to believe that venom exerts a decided action upon the capillaries themselves to bring about the remarkable ecchymoses found so commonly in cases of poisoning—an instance also of peripheral irritation, applicable here, is the effect of venom on the vagi peripheries in causing an in- creased respiration rate. ‘The ultimate fall of pressure seems to be cardiac in origin, since there is an accompanying diminution in the force of the beats. Section I1.—TuHe Action or VENOM GLOBULINS UPON THE BLooD PRESSURE. The Action of Venom Globulins upon the Blood Pressure of Normal Animals.— ‘Thirteen experiments were made with the globulins upon normal animals. ‘The doses usually given were those representing the amount of globulin in 0.015 gram of dried venom. ‘The results of all of these experiments indicate that all the globulins exert an action analogous to that of the pure venom, but that they exhibit a material difference in the relative degree of their toxicity. Of the thirteen experiments, seven were made with the water-venom-globulin, two with the copper-cenom-globulin, and four with the dialysis-venom-globulin. Of the first series, five were made with the globulin from the Crotalus adamanteus ; one with that of the Ancistrodon piscivorus, and one with that of the Cobra. ‘The second and third series were made with globulins from the Crotalus adamanteus venom. The water-venom-globulin produces the most profound changes, causing a primary diminution of pressure almost equalling that produced by pure venom, while dialysis-venom-globulin comes next; the copper-venom-globulin has~ but little effect. ‘The actions of all of these globulins is to cause a primary fall of pressure, which is followed by a rise towards the normal and more or less well marked, while if the dose is sufficiently large the rise is followed by a fall to zero at death. In one experiment made with the globulin from 0.0385 gram of dried Cobra venom there was no appreciable effect. This was probably due to the very small proportion of globulin in this variety of venom. Experiment No. 43. Time: Pressure REMARKS. min. sec. m. m. Normal as 110 Injected intravenously 0.0012 gram water-venom-globulin 10 80 (= 0.015 gram dried venom) from the dried venom of the 20 92 Crotalus adamanteus. 40 90 1 00 84 1 BO 84 3 00 95 5. 00 96 7 00 104 THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 103 Experiment No. 44. Normal Experiment No. 45. Normal Time: min, sec. 9 00 12 00 15 00 18 00 25 00 35 «6000 45 00 55 00 Time: min. sec. 10 20 40 1 00 1 20 1 40 3 40 5 40 7 40 9 40 10 00 10 20 10 40 14 00 liien0,0) 20 00 30 00 Time: min. Se Sa sec. 10 20 30 50 10 10 30 35 40 50 Pressure m. m. 106 106 110 116 124 130 130 130 Pressure m. m. 130 96 96 94 90 90 94 102 108 108 104 104 104 106 106 108 102 Pressure m. m. 120 86 96 86 90 84 90 80 104 100 82 REMARKS, Animal killed by pithing. Heart in diastole; some ecchy- moses in small intestine; blood remains fluid at the end of twenty-four hours—a few very soft clots are found. REMARKS. Injected intravenously the water-venom-globulin from 0.03 gram dried venom of the Crotalus adamanteus. Pressure falling. Clot formed in the canula. Injected water-venom-globulin from 0.015 gram dried venom. Animal killed by pithing. REMARKS. Injected intravenously 0.0033 gram water-venom-globulin (= 0.045 gram dried venom) from the dried venom of the Crotalus adamanteus dissolved by the addition of a trace of sodie carbonate. Injected 0.0066 gram as in the foregoing. Injected a similar dose. Killed by pithing. Heart arrested in diastole; few ecchy- moses; blood remains fluid after twenty-four hours. 104 Experiment No. 46. Normal Experiment No. 47. Normal Experiment No. 48. Normal THE VENOMS OF CERTAIN THANATOPHIDES. Time: min. sec. 10 20 30 50 1 00 i. BO 3 30 5 30 7 30 9 30 12 30 14 30 16 380 17 30 19 30 21 30 26 00 28 00 30 00 30 15 35 00 387 00 39 00 Time: min. sec. 30 50 100 1 30 i BO 2 00 2 40 3 10 Time: min. sec. 0 10 20 30 I a 2 30 5 30 10 30 14 30 19 30 Pressure / REMARKS. m. m. 148 Injected intravenously the water-venom-globulin from one 120 minim of fresh venom of the Crotalus adamanteus. 116 116 100 92 82 86 96 : 106 118 122 126 124 100 128 132 Clot in canula. 128 130 124 136 Clot in canula. 134 136 : 130 Animal killed by pithing; no ecchymoses; blood clots. Pressure REMARKS. m. m. 132 Injected intravenously the water-venom-globulin from 0.004 126 gram dried venom of the Ancistrodon piscivorus dissolved 124 in 1 ec. ec. distilled water by the addition of a few crystals of 136 sodic chloride. 122 114 Injected a similar dose. 106 116 104 Pressure REMARKS. m. m. 115 Injected intravenously the water-venom-globulin from 0.015 gram dried venom of the Crotalus adamanteus. 90 93 90 100 102 100 100 Hematuria. 103 THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 105 Time: Pressure REMARKS, min. sec. m. m. 24 30 95 29 30 85 34 30 80 42 30 75 - 47 30 73 52 30 60 af a 60 67 30 57 Ue. 8 55 80 30 38 85 ctor Dead; ecchymoses in intestines ; blood incoagulable. Experiment No. 49. Time: Pressure REMARKS. min, sec. m. m. Normal 6 O66 155 Injected intravenously water-venom-globulin from 0.035 gram 0 wants dried Cobra venom dissolved in 1 e. e. distilled water. 15 Bees 25 158 45 160 eld 158 2 00 157 4 00 153 8 00 153 13 00 153 18 00 143 23 00 153 c 28 Broke loose from canula. Experiment No. 50. Time: Pressure REMARKS. min. sec. m. m. Normal a) B% 126 Injected intravenously 0.0012 gram copper-venom-globulin 10 126 (= 0.015 gram dried venom) from the dried venom of the 20 126 Crotalus adamanteus. 30 132 50 126 2 50 124 4 50 126 6 50 126 & 60 124 10 50 124 Tit 5X0) 120 lv 20 110 18 20 114 Injected double the foregoing dose. 18 30 110 18 40 118 18) 50 112 19 00 112 20 00 120 22 00 116 2 one Killed. Heart in systole; few ecchymoses in lungs and intes- tines ; blood remains fluid after two hours 14 June, 1886. 106 Experiment No. 51. Normal Experiment No. 52. Normal THE VENOMS OF CERTAIN THANATOPHIDEA. Time: min. sec. 10 30 1 00 3 00 5 00 7 00 8 00 10 00 12 00 22 00 24 00 26 600 26 «10 26 30 26 40 27 00 27 ©6380 29 30 31 30 34 30 39 00 41 00 43 00 45 00 52 00 58 00 Time: min. sec. 20 40 50 1 20 3 20 5 20 18 20 18 23 18 30 18 45 OOS 19 25 1G) 55) 20° 25 20 55 Bl Bb) 22 00 30 00 Pressure m. m. 112 114 110 112 116 120 122 122 124 118 118 128 124 124 116 104 108 96 90 98 104 116 116 116 118 120 122 Pressure m. m. 132 124 112 116 108 108 120 130 96 100 94 102 96 76 60 46 42 REMARKS. Injected intravenously 0.0023 gram copper-venom-globulin = 0.03 gram dried venom) from the dried venom of the Crotalus adamanieus. Clot in canula. Injected double the dose. Killed by pithing ; no ecchymoses. REMARKS. Injected intravenously 0.0017 gram dialysis~venom-globulin from the dried venom of the Crotalus adamanteus dissolved in 1c. ec. distilled water with a trace of sodie carbonate. Tnjected 0.0034 gram dialysis-venom-globulin. Dead. No ecchymoses; heart in diastole; blood remains fluid at the end of one hour. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 107 Experiment No. 53. Time: min. sec. Normal 20 30 eacG 1 20 1 40 2 00 2 40 3 40 5 40 6 20 6 56 7 20 7 950 8 50 9 20 950) 10 50 11 50 12 50 14 20 14 50 15 20 15 50 Experiment No. 54. Time: min. sec. Normal : 10 20 30 1 00 3 00 8 OO fen 0) 16 00 Wy aX) 17 40 18 30 20 30 23 30 28 30 43 30 53 30 Pressure m. m. 126 120 114 110 102 102 100 -100 102 110 114 118 124 126 128 128 130. 154 122 112 112 84 78 62 Pressure m. m. 150 118 130 118 116 110 126 124 136 116 100 104 126 118 102 98 94 REMARKS. Injected intravenously dialysis-venom-globulin from the dried venom of the Crotalus adamanteus (quantity unknown). Injected more of the globulin. Killed by pithing. REMARKS. Injected intravenously 0.0017 gram dialysis-venom-globulin from the dried venom of the Crotalus adamanteus. Injected 0.0034 gram. “cc 4c Animal killed. The Action of Venom Globulins upon the Blood Pressure of Animals in which the Pneumogastric Nerves had been Severed.rtFour experiments were made on animals in which the pneumogastric nerves and depressor nerves were severed. The results in these experiments do not differ in quality from those obtained in 108 THE VENOMS OF CERTAIN THANATOPHIDEA. normal animals; the effects, however, appear to be less decided than in animals with the pneumogastrics intact. , Here, as in the previous experiments, the copper- venom-globulin exhibits comparatively little effect on the pressure. Of the four experiments which were made with globulins from the Crotalus adamanteus, one was made with the water-venom-giobulin ; two with the copper- venom-globulin, and one with the dialysis-cenom-globulin. It will be noticed that m several instances considerable rises of pressure occurred accompanied by struggles; the former effect being, no doubt, due to the latter, and not to a peculiar action of the globulin. Experiment No. 55. Time: Pressure REMARKS. min. sec. mm. m. Normal See 116 Injected intravenously 0.0011 gram water-venom-globulin from 10 100 the dried venom of the Crotalus adamanteus. 20 100 ; 30 110 i (0 106 1 40 110 3 40 110 5 40 108 Clot in canula. Animal killed by pithing. Experiment No. 56. Time: Pressure REMARKS. min. sec. m. m. Normal sista 130 Injected intravenously 0.0024 gram copper-venom-globulin 10 132 from the dried venom of the Crotalus adamanieus. 20 132 30 132 40 132 i @) 132 3 10 132 & 1 128 7 10 128 8 10 128 23 10 136 Injected a similar dose. 23 20 130 23 40 132 as s 23) 5D 132 24 00 116 Respiration greatly slowed. 24 15 116 Animal broke loose. Killed by pithing. Experiment No. 57. Time: Pressure REMARKS. min. sec. m. m. Normal 2 oe 116 Injected intravenously 0.0012 gram copper-venom-globulin 20 116 from the dried venom of the Crotalus adamanteus. 40 116 50 116 2°50 116 4 50 116 B HO. 116 8 50 112 THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 109 Time: min. 11 13 15 16 16 16 16 17 19 21 23 Experiment No. 58. sec. 50 50 50 10 20 30 45 45 45 45 45 45 45 00 Time: min. Normal or, 8 10 12 17 18 18 18 19 19 19 21 22, 23 25 27 29 34 34 38 41 47 49 sec. 10 20 30 50 50 SoowwseceoocoeocouarnwmrorRwWnwaw hy Ww wb SS a) Pressure m. m. 116 118 118 100 118 110 108 132 114 112 106 100 88 Pressure m. m. 120 100 112 110 150 190 130 120 126 122 120 118 118 100 148 140 170 176 144 140 166 122 128 114 82 80 60 50 28 REMARKS. Injected a double quantity. 6c iG ' 6 Struggles. Animal killed by pithing. REMARKS. Injected intravenously 0.0017 gram dialysis-venom-globulin from the dried venom of the Crotalus adamanteus. Struggles. Injected 0.0034 gram. Struggles. Injected a similar dose. “cc “ce ce Struggles. Dead. The Action of Venom Globulins upon the Blood Pressure of Animals in which the Pneumogastric, Depressor, and Sympathetic Nerves and Cervical Spinal Cord had been Cut.—Four experiments were made on animals in which the nerves of the 110 THE VENOMS OF CERTAIN THANATOPHIDEA. neck and the cord in the middle or upper cervical region (excepting one) were cut. They were all made with the globulins from the Crotalus adamanteus ; one with water-venom-globulin, one with 'copper-venom-globulin, and two with dialysis-venom- globulin. ‘The results of this series of experiments accord with those observed when pure venom was used, and with the preceding experiments with the globulins. ‘The primary fall of pressure is slight, while the tendency to a secondary rise is very marked, since in three of the experiments the pressure rose above the normal. The action of water-venom-globulin on the primary fall was most marked, while in the single experiment made with copper-venom-globulin, in which eight times the quantity was given in two doses, the pressure rose slightly, and continued above normal. When the dose is sufficient to kill, the pressure ultimately gradually declines, accompanied by a feeble pulse. In the last experiment with dialysis-venom-globulin it will be noticed that tremors are accompanied with a rise of pressure during their existence. Experiment No. 59. Time: Pressure REMARKS. min. sec. m. m. Normal Cnet 48 Section of cord made below the 6th cervical vertebra. Injected 10 38 intravenously 0.0011 gram water-venom-globulin from the : dried venom of the Crotalus adamanteus. 30 38 Artificial respiration stopped. 1 00 48 H 1@) 34 2 10 46 3 IO) 42 5 10) 40 it 10) 38 9 10 34 11 10 30 15 UO 30 17 40 30 19 OO 30 21 00 30 23 00 30 : 25 00 30 27 00 30 Animal killed by pithing. Experiment No. 60. Time: Pressure REMARKS. min. sec. m. m. Normal ele 29 Injected intravenously 0.0048 gram copper-venom-globulin 10 32 from the dried venom of the Crotalus adamanteus. 20 30 ; 40 32 OC) 32 3 30 38 5 30 40 7 30 42 THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 1]] Experiment No. 61. Normal Experiment No. 62. Normal Time: min. sec. 9 30 ll 30 13 30 16 30 17 +00 17 +30 18 00 20 00 24 00 26 00 28 00 30 00 32 00 34 00 Time: min. sec. 10 30 1 00 3 00 6 00 8 30 10 30 12 30 12 50 SO 13 30 13 50 15 00 16 00 18 00 Time: min. wu et ee Oo CO SD sec. 10 20 30 40 50 00 10 20 20 50 50 50 00 Pressure REMARKS. m. m. 42 42 40 44 42 Injected a similar quantity. 38 40 44 46 44 42 40 40 40 Animal killed by pithing. Pressure REMARKS. m. m. 42 Injected intravenously 0.0017 gram dialysis-venom-globulin 40 from the dried venom of the Crotalus adamanteus. 40 46 44 40 38 38 eae Injected 0.0068 gram. 44 48 46 48 36 46 28 Dead. Heart arrested in diastole; no ecchymoses; blood fluid. A few fibres of the anterior columns of the cord were uncut. Pressure REMARKS. m. m. 44 Injected intravenously 0.0068 gram dialysis-venom-globulin 40 from the dried venom of the Crotalus adamanteus. 40 52 Universal tremors persistent. 58 Clot in canula. 86 Blood pressure fell very low before this observation, and was raised by the tremors returning vigorously. Dead. No ecchymoses; blood incoagulable; heart natural, 112 THE VENOMS OF CERTAIN THANATOPHIDESA. From this series of experiments with globulins it seems clear that they possess the peculiar physiological effects of pure venoms upon the blood pressure; that the water-venom-globulin is the niost powerful, and the copper-venom-globulin the least so, and that the copper-venom-globulin seems to exhibit a more marked tendency than the others to cause a rise of pressure. Srotion I1].—Tue Action oF VENOM PEPTONES UPON THE BLOOD PRESSURE. The Action of Venom Peptones upon the Blood Pressure of Normal Animals.— Seven experiments were made with the peptones from different venoms: two with that of Crotalus adamanteus ; three with Ancistrodon piscivorus; and two with Cobra. ‘The action of peptones upon the blood pressure is similar to that observed with the pure venom and the globulins, but their power to cause the primary pro- found fall of pressure is certainly much less, while the rise of pressure after the primary fall is decidedly more marked, and there is also a tendency to go above the normal, In two experiments, one with the peptone of the Crotalus and one with that of the Moccasin, the pressure was not primarily reduced, but there was a rise above the normal from the first. Where the animal was watched until death the pressure was observed to undergo a more or less gradual decline with feeble heart- beats. In several instances a rise of pressure was noted which was usually due to convulsive seizures. Experiment No. 63. Time: Pressure REMARKS. min. sec. m. m. Normal ase 140 Injected intravenously the peptone from 0.015 gram dried 10 128 venom of the Crotalus adamanteus. 20 128 30 136 40 128 50 128 00 124 11 00 116 21 00 116 Clot. 49 aoa Dead. No ecchymoses; lungs seem congested; blood clots readily. : Experiment No. 64. Time: Pressure REMARKS. mim. sec. m. m. Normal 8) fe 114 Injected intravenously the peptone from 0.03 gram dried 10 130 venom of the Crotalus adamanteus. 30 132 00 120 2 00 140 5 00 144 9 00 124 Killed. Hechymoses in the lungs; blood clots. THE ACTION OF VENOMS UPON ARTERIAL PRESSURE. 113 Experiment No. 65. Normal Experiment No. 66. Normal Experiment No. 67. Normal 15 Time: min. sec. pee 10 10 1k 11 30 00 30 30 30 50 10 30 Time: min, sec. OTT OD OF Oo Oo bo WP ee 10 20 30 00 30 00 30 00 00 20 40 00 30 00 30 00 Time: min. aor op bP we He co Ss 9 13 June, 1886. sec. 10 20 30 40 50 00 20 40 00 20 40 40) 50 00 20 30 30 30 Pressure m. m. 86 88 88 88 92 94 100 102 96 Pressure m. m. 116 94 66 70 76 74 76 76 76 66 60 66 68 66 62 56 60 Pressure m. m. 140 94 100 160 170 190 130 130 142 136 136 136 118 114 104 112 112 118 122 REMARKS. Injected intravenously the peplone from 0.015 gram dried venom of the Ancistrodon piscivorus. Injected double the amount. REMARKS, Injected intravenously the peptone from 0.015 gram dried venom of the Ancistrodon piscivorus. Injected a similar quantity. Injected a double quantity. REMARKS. Injected intravenously the peptone from 0.05 gram dried venom of the Ancistrodon piscivorus. Convulsions. Injected 0.005 peptone. Killed. 114 THE VENOMS OF CERTAIN THANATOPHIDES. Experiment No. 68. Time: Pressure REMARKS. min. sec. m. m. 2 Normal alee 140 Injected intravenously the peptone from 0.005 gram dricd 10 130 Cobra venom. 20 148 30 140 40 142 1 00 142 1 20 140 3 20. 138 6 20 152 Il 20 224 Convulsive twitchings. 16 20 176 16 50 134 Wie x0 92 18 20 104 Blood is asphyxiated ; no respiration. 18 50 84 19 00 46 1G) IO 46 1930 38 19 50 36 20 10 40 20 30 40 Killed. Experiment No. 69. Time: Pressure REMARKS. min. sec. m. m. Normal ascites 130 Injected intravenously the peptone from 0.06 gram dried 10 130 Cobra venom. 15 116 30 156 50 156 3 20 140 8 20 190 Tonic convulsions. 10 20 176 Convulsive twitchings; asphyxiated blood; respiration 10 30 208 ceased. 10 40 182 10 50 136 1l 00 116 il QO 102 11 40 86 12 00 66 12 20 54 12 40 38 Dead. The Action of Venom Peptones on the Blood Pressure of Animals with Pnewmo- gastric and Depressor Nerves Severed.—After section of the pneumogastric and depressor nerves the results are not appreciably altered. Four experiments were made: one with Crotalus adamanteus ; one with Ancistrodon piscivorus, and two with Cobra venom. In all of these experiments the pressure during the secondary rise went above the normal. THE ACTION Experiment No. 70. Normal Experiment No. 71. Normal Experiment No. 72. Normal Time: min. sec. or 00 OD et et et 20 25 9 9) 35 40 45 50 62 10 15 30 40 50 00 20 30 30 30 30 30 30 30 00 00 00 00 00 Time: min. sec. Jy RO) JRO) 1K) PEN J Hm Co Co CO 10 20 30 40 50 00 20 » 40 00 30 40 90 00 10 20) 20 Time: min. sec. 10 20 40 00 30 OF VENOMS UPON ARTERIAL PRESSURE. 115 Pressure REMARKS. m. m. 160 Injected intravenously the peptone from 0.015 gram dried 160 venom of the Crotalus adamanteus. 140 150 156 Struggles. 156 134 158 164 126 122 Struggles. 146 148 140 140 136 148 142 140 138 Killed. Pressure REMARKS. m. m. 124 Injected intravenously the peptone from 0.015 gram dried 100 venom of the Ancistrodon piscivorus. 140 _ 150 130 140 134 114 116 136 124 110 124 136 110 96 Respiration ceased; heart beats. Pressure REMARKS. m. m. 138 Injected intravenously the peptone from 0.005 gram dried 142 Cobra venom. 136 134 140 138 116 THE VENOMS OF CERTAIN THANATOPHIDEA. Time: Pressure REMARKS. min. sec. m. m. 3 30 136 5 30 136 7 30 132 9 B80 132 10 00 140 21 30 146 Twitchings. 25 30 142 26 30 150 Killed. No ecchymoses. Experiment No. 73. Time: Pressure REMARKS. min. sec. m,. m. S Normal Pars 124 Injected intravenously the peptone from 0.006 gram dried 10 122 Cobra venom. 20 128 40 124 1 00 126 3 00 120 5 00 120 16 00 118 34 00 Soe Dead. Asphyxiated; no ecchymoses; blood clots in canula. The Action of Venom Peptones cn the Blood Pressure of Animals in which the Pneumogastric, Depressor, and Sympathetic Nerves and Cervical Spinal Cord were Cut.—In five experiments on animals in which the nerves in the neck and the spinal cord in the middle or upper cervical region were cut we found that but little alteration occurred in the blood pressure until late in the poisoning, excepting im one experiment with the Ancistrodon piscivorus, im which the pressure sunk imme- diately and death occurred in thirty seconds. ‘Two experiments were made with the peptone of the Crotalus adamanteus ; one with the Ancistrodon piscivorus, and two with the Cobra. In all of these experiments, excepting one with Cobra, there was an immediate comparatively slight fall of pressure after injection, which was followed generally by a rise; in the excepted case of the Cobra there was a primary rise equal to 3 m.m. of mercury, which was followed by a fall, and this in turn by arise. The pressure, as in the previous experiments, usually declines towards death. Experiment No. 74. Time: Pressure REMARKS. min. sec. m. m. Normal Badia 50 Tnjected intravenously the peptone from 0.015 gram dried 10 50 venom of the Crotalus adamanteus. 20 48 4() 48 1 00 48 3 00 44 6 00 42 11 00 49 13 00 42 15 00 42 THE ACTION Time: min, sec. Uy OO 17 30 19 30 21 30 23 30 25 30 27 «630 29 30 31 30 34 30 OF VENOMS UPON ARTERIAL PRESSURE. 117 Pressure m. m. 50 50 48 48 46 46 44 44 42 38 Experiment No. 79. Time: Normal min. — bo sec. 15 20 30 00 30 00 Pressure mM. m. 67 50 50 50) 50 50 00 Experiment No. 76. Normal Time: min. sec. 10 18 30 1 00 Pressure m.m, 50 45 38 33 Experiment No. 17. Normal Time: min. sec. 10 20 30 1 00 12 00 12 30 30 Pressure m. m. 128 122 132 134 132 138 136 eee REMARKS. Injected the peptone from 0.03 gram dried venom. Dead. No ecchymoses; blood fluid after fifteen minutes. REMARKS. Injected intravenously the peptone from 0.015 gram dried venom of the Crotalus adamanteus. REMARKS, Injected intravenously the peptone from 0.015 gram dried venom of the Ancistrodon piscivorus. Dead. REMARKS, Injected intravenously the peptone from 0.01 gram dried Cobra venom. Dead. Blood clots readily. Ecchymoses in base of lungs. 118 THE VENOMS OF CERTAIN THANATOPHIDESA. Experiment No. 78. Time: Pressure ~ REMARKS. min. sec, m,. m. Normal he 38 Injected intravenously the peptone from 0.015 gram dried ll 39 Cobra venom. 20 41 40 40 5 1 40 35 5 40 35 10 40 37 15 40 38 16 00 okt F ee ae aR 28 t Injected a similar dose. LGU 38 16 25 43 16 45 43 17 00 43 18 00 40 20 00 25 23 00 sa Dead. From all of the results of these experiments it seems justifiable to conclude that the isolated principles of venoms exert the poisonous actions of pure venoms on the blood pressure, and that their toxic effects are essentially simply different in degree. ‘These various poisons all play a part in the alterations of pressure, acting towards the same end, but mainly with different degrees of intensity; the ewater- venom-globulin appears to be the most potent in the pressure alterations, the dialy- sis-venom-globulin next, then the peptone, and finally the copper-venom-globulin. The globulins are the more active in the production of the diminution of pressure, and the peptone in the secondary rise. The globulins no doubt play a very important part in the poisonous phenomena of Crotalus poisoning, a less important part in Ancis/rodon poisoning, and but very little in Cobra poisoning; these differences not depending as much upon differences in the quality of the globulins in the species of venom to which they belong as on differences in quantity. THE ACTION-OF VENOMS UPON RESPIRATION. 119 (lal e P AE Ik IOC THE ACTION OF VENOMS AND THEIR ISOLATED GLOBULINS AND PEPTONES UPON RESPIRATION. Srection I.—PurE VENOM. In our experiments on respiration rabbits were always used, and the rate of breathing was recorded on a revolving drum by the lever of a Marey’s tambour, the latter being connected with the animal by means of a tracheal tube. The injections in all of the experiments, excepting two, which were subcutaneous, were made into the external jugular vein. ‘In experiments on normal animals we observed no qualitative difference in the several venoms used. ‘Ten experiments were made upon normal animals: four with the venom of the Crotalus adamanteus; three with that of the Moccasin, piscivorus, and three with that of the Cobra. In eight of these experiments there was a primary increase in the respiration rate followed by a diminution far below the normal, while m two the respirations were at once diminished, and became per- sistently slower until death. In both of these cases death occurred very soon after injection, indicating a most profound action of the poison. . Action of the Pure Venoms on the Respirations in Normal Animals. Experiment No. 1. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m,. Normal ... 84 10 Injected intravenously 0.002 gram dried venom of the 10 180 16 Crotalus adamanteus dissolved in 1 e. ¢. distilled 40 84 12 water. 1 96 2 20 108 O00 4 50 72 8 6 50 72 6 8 50 60 6 10 50 48 ates Convulsive movements. Il 20 40 4 te fs 11 50 24 4 “s « 12 20 26 12 50 eA are Dead. 120 THE VENOMS OF CERTAIN THANATOPHIDESA. Experiment No, 2. Time: min. Normal oo to) b> sec. 10 40 00 10 40 00 10 Respirations per minute. 42 43 84 30 10 Experiment No. 8. Time: min. Normal m 0) co Do wD ee Coreen sec. 40 00 30 00 30 00 30 30 30 30 30 Respirations per minute. 84 158 120 96 90 96 102 120 35 60 35 10 Experiment No. 4. Time: min. sec. Normal b Ww ee 15 2 oO 00 50 20 25 Respirations per minute. 66 Length of / curve m.m. 6 10 12 25 23 14 Length of curve m.m. 7 3 7 11 10 12 10 7 5 6 Pr i) Length of eurye m.m. 6 16 16 6 5 REMARKS. Injected intravenously 0.004 gram dried venom of the Crotalus adamanteus dissolved in 1 ec. ¢. distilled water. Struggles, which prevent a count. Convulsive movements. Conjunctival reflexes gone. Respiration ceased. Heart still beating. The respira- tory muscles respond to stimulus. The spinal cord was exposed, and the motor columns were found to respond to electrical stimulus. The motor nerves responded after the motor columns of the cord had lost their irritability. REMARKS. Injected intravenously 0.006 gram dried venom of the Crotalus adamanteus dissolved in 3 minims distilled water. Strugeles. Conjunctival reflexes gone. Respiration ceased. Respiratory muscles irritable. The spinal cord was quickly exposed; the sensory columns give no response, the motor columns are active. The motor columns of the cord fail before the motor nerves. REMARKS. Injected intravenously 0.015 gram dried venom of the Crotalus adamanteus dissolved in 1 ec. ec. distilled _ water. Arrest of respiration attended with a tetanic condition. Respiration ceased. Spinal cord rapidly exposed and tested by electrical currents; sensory columns fail first, then the motor columns, then motor nerves. THE ACTION OF VENOMS UPON RESPIRATION. 121 Experiment No. 5. Normal Time: Respirations min. sec. per minute. Sasi 100 10 210 20 150 30 140 40 120 50 500 eo 30 Experiment No. 6. Time: Respirations min. sec per minute, Normal O 6 135 10 420 20 270 30 65 40 60 50 120 0,0 120 i u@ 60 1 20 90 6' 20 60 11 20 180 16 20 210 16 30 Experiment No. 7. Time: Respirations min. sec. per minute, Normal ... 144 10 300 20 240 30 150 5 00 60 7 00 80 12 00 54 18 00 70 18 05 ee: 18 30 210 18 40 160 18 50 80 23 50 65 16 June, 1886. Length of curve REMARKS. m.m. 9 Injected intravenously 0.004 gram dried venom of the 8 Ancistrodon piscivorus dissolved in 5 minims dis- 21 tilled water. 20 23 508 Convulsions. 000 Dead. Length of curve REMARKS. m.m. 6 Injected intravenously 0.004 gram dried venom of the Ancistrodon piscivorus dissolved in 1 ¢. ¢. distilled water. Struggles. Respiration at once began to increase 6 rapidly, and reached a maximum rapidity during the occurrence of struggles. 18 Tetanic movements. Bee “cc “cc 10 “ec oo 12 ie Killed. Length of curve REMARKS. m. m. 6 Injected intravenously 0.004 gram dried venom of the 8 Ancistrodon piscivorus dissolved in 1 ¢. ¢. distilled 12 water. 11 10 7 7 1 Injected as above 0.008 gram venom. ©! 10 7 Killed by pithing. 122 THE VENOMS OF CERTAIN THANATOPHIDE A. Experiment No. 8. Length of 5 Time: Respirations ‘ curve REMARKS. min. sec. per minute. m.m. Normale 60 9 Injected intravenously 0.015 gram dried Cobra venom 20 80 eis dissolved in 1 ¢. ¢. distilled water and filtered. 40 120 15 Struggles. 1 00 100 45 is 1 20 60 10 1 40 42 38 2 00 C00 Bees Respiration ceased. Experiment No. 9. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m. Norma ene 300 8 Injected intravenously 0.015 gram dried Cobra venom 10 300 9 dissolved in 1 ¢. c. distilled water. 20 255 11 30 285 10 40 240 10 1 00 255 11 1 30 200 i 2 00 150 iv 2 10 125 7 8 20 Roar evs Respiration ceased. Experiment No. 10. Time: Respirations REMARKS. min. sec. per minute. NOME 5 6 « 36 Tnjected intravenously 0.003 gram dried Cobra venom in solution. 20 39 40 39 1 00 51 1 30 52 2 00 45 4 00 48 9 00 46 12 00 36 14 Respiration ceased. The Action of Pure Venoms on the Respiration in Animals in which the Pneu- mogastric Nerves were Cut.— When injections are made, after section of the pneu- mogastric nerves, the primary increase in the respiration rate does not occur, but a diminution begins at once; and, on the whole, drops irregularly until death ensues. Four experiments were thus made: one with Crotalus adamanteus, two with Ancistrodon piscivorus, and one with Cobra venom; the results being on the whole reasonably uniform. THE ACTION OF VENOMS UPON RESPIRATION. 123 Experiment No. 11. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m. Normal 42 7 Injected intravenously 0.002 gram dried venom of the Crotalus adamanteus dissolved in 1 ec. c. distilled water. 30 20 18 Slight struggles preceding this observation interfered 1 00 28 10 with the marker. 1 30 6 22 2 00 6 19 2 3 3 25 3 00 12 18 3 30 6 21 4 3 6 21 5 30 6 15 6 30 8 12 Experiment No. 12. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m. Normal 103 15 Injected intravenously 0.004 gram dried venom of the 10 94 15 Ancistrodon piscivorus dissolved in 1 ec. ¢. distilled water. 20 84 30 Struggles. : 30 102 25 40 17 25 50 60 28 1 00 60 25 eo 45 30 It 15) a06 Dead. Respiration ceased; heart still beats. Animal dies in tetanus. Experiment No. 13. Time: Respirations REMARKS. min. sec. per minute. Normal 102 Injected intravenously 0.003 gram dried Cobra venom dissolved 30 57 in | ¢. ¢. distilled water. 1 00 78 2 00 oT 4 00 66 10 soe Experiment No. 14. : Length of Time: Respirations curve REMARKS. min. sec. per minute, m.m. Normal 127 17 Injected intravenously 0.004 gram dried venom of the Ancistrodon piscivorus dissolved in 1 ¢. ec. distilled water. 20 92 47 30 90 32 40 82 50 82 05 67 1 30 wets Respiration ceased. 124 ' THE VENOMS OF CERTAIN THANATOPHIDEA. In none of these experiments do we find a primary increase in the respiration rate, as in animals with intact vagi, but invariably a diminution. It seems clear, therefore, that the first result must be dependent upon an excitation of the peri- pheries of the pneumogastric nerves, and that the diminution of respirations is due to acentrally active cause. Should the lessened number of the respirations be central, that is, dependent upon a depression of the respiratory centres, we would expect to find that the degree of depression would depend upon the relative amount of venom coming in contact with these centres in a given space of time. We have accordingly made an experiment, in which this suggestion is admirably carried out by injecting the venom into the carotid artery, thus throwing the poison directly upon the respiratory centres. Experiment No, 15. Length of Time: Respirations curve REMARKS, min. sec. per minute. m.m. : NORMA 6 5 - 18 42 Injected into the right carotid artery 0.015 gram of 15 1 38 dried venom of Crotalus adamanteus dissolved in 1 30 4 30 ec. c. distilled water. 1 00 4 25 I ait) 10 45 Convulsions. 2 00 tore haps Dead. It seems obvious from the preceding experiments that venoms exert a double action on the respiration; first, an irritant action on the peripheries of the pneu- mogastric nerves, by which an increase in the respiration rate is brought about; and secondly, a depression of the respiratory centres, by which the respiration rate is diminished. Since the diminution in the respirations occurs in animals with cut pneumogastrics immediately after injection, and at a time when an increase occurs in normal animals, it is apparent that these two factors are acting in normal animals at the same time to produce opposite results; consequently, whether we have an increase or a decrease in the respirations must be dependent upon the relative degree of power exerted by one or the other of these factors. In most cases we have found a primary increase of respirations followed by a diminution ; it is therefore obvious that the action of the venom upon the peripheries of the pneumogastric nerves was more than able to compensate for the depressant action of the poison upon the respiratory centres; this is very clear since no increase of respirations above normal occurs in animals with cut pneumogastrics. In the two cases in normal animals in which a decline from the first was observed, and in which the animals died in a few minutes after injection, the action of the venom upon the respiratory centres was so profound that the accelerator factor was unable to cause arise. ‘This is also illustrated in the experiment in which the venom was injected into the carotid artery and thrown upon the respiratory centres. Since venom does not seem to exert other than a depressant action upon the respiratory centres, it does not appear probable that it would have an opposite effect upon the respiratory nerves, so that the effect of the venom upon the peri- pheries of the pneumogastric nerves is probably one of irritation rather than stimu- THE ACTION OF VENOMS UPON RESPIRATION. 125 lation, and probably due to some secondary cause, which is likely to be located in the profound alteration of the blood or the destructive action of the venom upon the pulmonary tissues, as illustrated, for instance, upon capillaries. Section IJ.—Tur Action of GLOBULINS ON THE RESPIRATIONS. The Action of Venom Gtlobulins upon the Respiration in Normal Animals.— Seven experiments were made with globulins upon normal animals: three with the water-venom-globulin of the Crotalus adamanteus ; and one with the water-venom- globulin of Cobra; one with the copper-venom-globulin, and one with dialysis-venom- globulin, both from the Crotalus adamanteus. ‘These poisons, excepting the copper-venom-globulin, all act like the pure venoms, but generally with a less degree of intensity, causing a primary acceleration of the respiration followed by a decline. In the second experiment, however, there was no diminution, but the respirations became enormously increased so that at death they were nearly trebled in frequency. ‘The copper-venom-globulin does not cause any primary acceleration, but simply a diminution. Experiment No. 16. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m. Normale = = - 100 9 Injected intravenously the water-venom-globulin from 20° 100 15 0.015 gram dried venom of the Crotalus adamanteus. 40 100 12 1 90 96 11 3 00 96 9 4 00 120 10 5 00 120 10 6 00 132 10 8 00 100 10 10 00 90 9 12 00 80 9 14 00 69 7 14 05 me act Injected as above from 0.06 gram dried venom. 14 20 80 15 14 40 60 10 Struggles. 16 40 90 10 18 40 96 8 20 40 108 9 22 40 114 9 24 40 108 9 26 40 90 6 28 40 96 8 30 «40 72 vf 32 40 64 8 34 40 70 8 36 40 75 8 38 40 80 9 40 40 90 10 44 00 300 apie Dead. Heart arrested in diastole ; some ecchymoses. 126 THE VENOMS Experiment No. 17. Normal Time: min. sec. 15 30 1 00 1.0 6 30 Ik a) 16 30 17 00 27 00 29 00 Respirations per minute. 19 80 60 60 90 110 110 110 120 190 Experiment No. 18. Normal Time : min, sec. 15 25 45 I i16 2 00 6 00 IO. 16 00 26 00 33 «(00 56 00 66 00 76 00 120 Respirations per minute. 114 126 132 150 150 204 114 84 62 60 Experiment No. 19. Normal Time: min. sec. 20 30 0 2 30 5730 10 30 14 30° 19 30 24 30 29 30 34 30 42 30 47 30 52 30 Respirations per minute. 13 84 i. 120 108 96 96 82 70 15 78 72 69 12 96 OF CERTAIN THANATOPHIDESA, Length of » curve REMARKS. m, mM. 8 Injected intravenously 0.0158 gram water-venom-globuc lin (5 days old) from the dried venom of Crotalus adamanteus. 8 Struggles. Injected the same as above. Dead. Blood remains fluid; some ecechymoses. REMARKS. Injected intravenously the water-venom-globulin from 0.035 gram of dried Cobra venom dissolved in 1 ¢. ec. distilled water. Killed. Animal in fair condition. REMARKS. Injected intravenously the water-venom-globulin from 0.015 gram dried venom of the Crotalus adamanteus. Hematuria. THE ACTION OF VENOMS UPON RESPIRATION. D7 Time: Respirations REMARKS. min. sec. per minute. 57 30 90 67 30 : 84 i 84 80 30 12 8) ae Dead, Hechymoses generally ; blood fluid. Experiment No. 20) Length of Time : Respirations curve REMARKS. min. sec. per minute. m.m. Normal .. . 180 20 Injected intravenously the copper-venom-globulin from 20 174 19 0.015 gram dried venom of the Crotalus adamanteus. 40 168 19 C0 168 19 3 30 168 20 5 30 108 15 7 30 100 9 9 30 110 10 IL 00 168 17 3 00 138 11 15 00 120 10 17 00 144 10 19 00 102 9 21 00 108 10 23 00 104 T 25 00 112 9 27 00 100 9 30 00 90 7 34 00 90 7 39 00 112 10 41 00 85 8. 43 00 96 7 45 00 108 b) 47 00 108 7 49 00 76 6 51 00 66 7 53 00 80 8 57 00 96 8 59) +00 90 9 60 00 116 10 63 00 118 8 65 00 116 10 Struggles. 69 00 104 9 00 100 7 73 00 116 7 75 00 100 8 77 «00 116 11 79 00 140 1] 81 00 130 10 85 00 130 10 87 00 120 10 91 00 126 9 92 00 PreK alee Killed by pithing. Lungs very much ecchymosed; abdo- minal viscera normal; heart normal; blood coagulates. 128 THE VENOMS OF CERTAIN THANATOPHIDEA. Experiment No. 21. Normal Time: min. sec. TO Ot Re 8 10 11 13 15 17 17 17 18 19 21 24 27 27 28 29 10 26 40 00 00 00 00 00 00 30 oo So oOnwmrrnwnnwnbd wo Ww WH COC Se Seeeoqeo Ee © Oo 2S Se Respirations per minute. 54 60 o4 54 48 60 72 54 63 60 60 72 70 78 78 102 72 66 60 70 60 60 60 Experiment No. 22. Normal Time: min. sec. bo 10 12 13 13 14 19 24 29 Q t) 54 59 10 20 30 40 00 00 00 00 00 30 50 30 00 00 00 00 00 00 Respirations per minute. 112 120 160 140 140 140 126 156 174 130 142 150 132 130 120 110 80 80 Length of / curve REMARKS m.m. 18 Injected intravenously 0.0012 gram water-venom- 16 globulin from the dried venom of the Crotalus ada- 15 manteus. 15 16 17-42 Struggles. 33 30 32 28 30 42 38 45 Injected 0.0022 gram water-venom-globulin. 42° 38 38-78 Struggles. 22 23 25 29 Injected 0.0024 gram water-venom-globulin. 26 ' 25 Killed by pithing ; some ecchymoses. Length of i curve REMARKS m.m. 9 Injected intravenously the dialysis-venom-globulin from 12 0.015 gram dried venom of the Crotalus adamanteus. ° 16 16 15 16 10 14 15 14 10 Injected dialysis-venom-globulin from 0.06 gram of 22 dried venom. 13 Dead. Respiration ceased before the heart. Hechy- moses in the lungs and in the pericardium, in the small intestine, ureters, and bladder. THE ACTION OF VENOMS UPON RESPIRATION. 129 The Action of Venom Globulins on the Respiration of Animals in which the Pneumogastric Nerves were Cut.—Iwo experiments were made on animals with cut pneumogastric nerves: one with the dialysis-venom-globulin, and one with the copper-venom-globulin, both from the Crotalus adamanteus. In neither experiment was there an increase in the respirations; these results being in accord with the experiments made with pure venom. Eaperiment No. 23. Time: Respirations min. sec. per minute. Normal : 42 10 39 20 30 40 24 1 00 Py ( 1 20 ? 3 20 20 5 20 35 8 20 24 18 20 30 23 20 32 36 20 42 36 40 42 38 40 24 42 00 Eaperiment No. 24. Time: Respirations min. sec. per minute. Normal 60 30 54 1 00 48 3 00 48 8 00 48 13 00 52 15 00 48 15 30 30 15 40 54 16 00 ? 16 30 18 19 00. 12 24 00 20 27 00 20 30 00 30 35 00 26 39 00 27 41 00 30 44 00 42 49 00 30 54 00 17 June, 1886, Length of curve m.m. 8 Length of curve B TE DAD ANH a on , — . hte AarrnrngopRe. REMARKS. Pneumogastrie nerves previously cut. Injected hypo- dermically the dialysis-venom-globulin from 0.015 gram dried venom of the Crolalus adamanteus. Struggles. Injected dialysis-venom-globulin from 0.06 gram.' Struggles. Respiration ceased; heart beats feebly; blood remains incoagulable; great ecchymoses in abdominal viscera. REMARKS. Pnenmogastric nerves previously cut. Injected intra- venously the copper-venom-globulin from 0.015 gram dried venom of the Crotalus adamanieus. Injected copper-venom-globulin from 0.03 gram dried venom in two doses. Struggles with very irregular breathing followed by gasping respiration. Injected copper-venom-globulin from 0.12 gram dried venom in two doses. Respiration ceased ; heart still beats; ecchymoses in heart and lungs marked. 130 THE VENOMS OF CERTAIN THANATOPHIDES. The results of these experiments with the globulins indicate that the water- venom-globulin and dialysis-venom-globulin act like the pure venom, while the copper-venom-globulin lacks the property of producing the primary acceleration of the respirations. Section IIJ.—Tur Action or VENOM PEPTONES ON THE RESPIRATION. The Action of Venom Peptones on the Respiration in Normal Animals.—Three experiments were made on the normal animals with the venom peptones; in two with the peptone from the Crotalus adamanteus, and in one with the peptone from the Ancistrodon piscivorus. In all of these experiments the increase of the respi- ration rate was strongly marked. Experiment No. 25. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m. Normale sear 225 68 Injected intravenously the peptone from 0.03 gram 10 255 60 dried venom of the Crotalus adamanteus obtained 30 255 60 by boiling. 1 00 300 56 2 OO . 270 50 5 00 270 50 9 00 270 55 Killed. Blood clots readily; moderate ecchymoses in the lungs Experiment No, 26. Length of Time: Respirations curve REMARKS. min. sec. per minute. m.m. NWomml 5 o « 180 11 Injected intravenously the peptone from 0.06 gram 10 240 16 dried venom of the Ancistrodon piscivorus obtained 30 270 15 by boiling. 40 240 he 1 00 345 1 1@ 270 20 240 4 20 240 S20 300 a 18° 20 360 14 28 20 270 38 20 180 ll THE ACTION OF VENOMS UPON RESPIRATION. 1B Experiment No. 27. Length of Time: Respirations curve REMARKS. min. sec. per minute, m. m. Normal ... 75 9 Injected intravenously the peptone from 0.015 gram 10 120 42 dried venom of the Crotalus adamanteus. 3 00 30 10 6 00 75 8 11 00 50 8 18 00 48 7 23 «00 50 7 28 00 45 it oe 00 60 7 ar (Oo) 60 9 49 S00 bine Dead. No ecchymoses; lungs slightly congested. In one animal the increase was equal to one-third of the normal; in the second, in which a larger dose was used, the normal rate was doubled; and in the third it rose to more than one-half of the normal. There was not, however, in any of the animals that marked depression which is observed in poisoning with pure venom or venom globulins, The Action of Venom Peptones on the Respiration in Animals in which the Pneumogastric Nerves had been previously Divided.—In one experiment in which the pneumogastric nerves were cut and in which the peptone from the venom of the Crotalus adamanteus was used, the well-marked primary increase in the respi- rations did not occur, there being a diminution from the first. Experiment No. 28. Length of Time: Respirations curve : REMARKS. min. sec. per minute. m.m. Nonmale ne. 80 13 Injected intravenously the peptone from 0.015 gram 10 52 18 dried venom of the Crotalus adamanteus. 15 37 12 20 25 8 30 22 7 1 00 30 8 i & 30 7 3 30 40 9 8 30 60 8 Struggles. 15 30 36 13 20 30 48 17 25 30 50 14 30 30 52 15 35 «(00 55 12 40 00 48 15 45 00 45 15 50 00 45 16 62 00 44 15 Killed by pithing. In this experiment, as in those with pure venom and venom globulins in which the animals had the pneumogastrics cut, the increased respiration rate seen in normal animals did not occur. 132 THE VENOMS OF CERTAIN THANATOPHIDESA. The results of the experiments with venom peptone are therefore in accord with those with the pure venom and. the venom globulins. . Summary.—From the results of the observations with pure venoms and their globulins and peptones upon the respiration it seems clear that the primary action of all of the above poisons, excepting the copper-venom-globulin, is to cause an increase in the number of respirations, and secondarily to diminish the respirations below the normal. Of the different principles the peptone seems to exert the most decided power in causing the acceleration, while the copper-venom-globulin seems to utterly lack this action. Since the primary increase of the respirations does not occur in any case after section of the pneumogastric nerves, this effect must be exerted by an action of the poisons upon the peripheries of these nerves, and since after section of these nerves a diminution of the respirations always occurs this effect must be due to a depres- sion of the respiratory centres, as we have found that the motor nerves and muscles of respiration are irritable long after the cessation of this function. PATHOLOGY. 133 CREIPAG eee hi Reo eNe. PATHOLOGY. Pathology of Serpent Venoms.—The pathology of snake poisoning in man owes most of what is best in our knowledge of it to the researches of the East Indian surgeons and to American observers. In the following observations Prof. H. F. Formad has followed with great suc- cess the lines of a research which were laid down with care by the authors of this essay. They have also been at great pains to repeat, and to verify, most of the observations made by this distinguished observer. The Nature and Character of the Individual Morphological Constituents of Venom.—Having seen that fresh venom consists morphologically of a liquid and of a solid part, it was necessary to ascertain the exact nature and character of each. The following means were resorted to :— Ist. The separation of the granular material (of fresh venom) by filtration and the submission to physiological tests of the liquid filtrate and of the solid residue, each separately. ; 2d. ‘The exposure of fresh venom to a temperature high enough to kill organized life, and then submitting it to physiological tests. 3d. Studying the effects of venom and of its isolated morphological constituents upon dead animal substances. (Putrefaction and other experiments.) 4th. The isolation and culture of the organisms contained in venom and the testing of the physiological effects of these isolated and washed organisms (viz., of pure cultures of micrococci). Ist. Filtration Experiments with Fresh Venom.—On account of its viscid and glutinous character venom could not be satisfactorily filtered except under a high pressure through a vacuum filter. About two drachms of fresh Crotalus adamanteus venom were forced by means of a hydraulic air pump through a porous clay cylin- der such as is employed in certain small galvanic batteries, or else the venom was filtered through a thin layer of plaster of Paris moulded in the neck of a small glass filter. The liquid filtrate obtained was perfectly clear, and examined under the microscope showed no organic or solid particles of any kind. ‘The solid residue left upon the filter consisted of granular material, such as has been described before, of bacteria and a few cells. This residue was diligently and repeatedly washed with boiled distilled water, by passing the latter through the filter. The amount of residue (about three grains) just obtained was dried and intro- duced subcutaneously into the pectoral muscle of a pigeon, but without effect. 134 THE VENOMS OF CERTAIN THANATOPHIDES. Two pigeons were injected in the pectoral muscle, one with five, and the other with two minims of the liquid filtrate above described, and both died promptly within six minutes and twenty-five minutes respectively. 2d. Experiments with Heated Venom.—Fresh Crotalus venom rapidly dried was put in a covered watch-glass and subjected for one hour to a temperature of 115° C. in the dry-heat oven. ‘The venom was thereby converted into a dense resinous opaque brown mass, Two grains of this mass, upon the addition of distilled water forming a turbid liquid, were divided into thirds and injected hypodermatically into a rabbit, a rat, and a pigeon, respectively. The rabbit died in 15 minutes, the rat in 12 minutes, and the pigeon in 7 minutes, after the operation, with results and lesion similar to those obtained by the use of fresh venom. This experiment also shows that the virulence of venom does not reside in any of its organized constituents. 3d. Putrefaction Eaperiments.—The testing of the effects of venom on various dead animal substances was particularly desirable on account of the remarkable capacity of the venom to induce rapid putrefaction in the tissues of living animals. It was necessary to learn whether this property of bringing about speedy necrotic changes was an action inherent in venom or due to any of its accidental constituents. Putrefaction Experiments with Sterilized Bouillon and Fresh Venom and its Active Principles (not Sterilized).—This bouillon was prepared from chicken in the same manner as that ordinarily used for culture liquids for bacteria, and the experiments were executed in a room at a temperature of about 70° F. About two drachms of sterilized bouillon were put in each of sixteen ordinary test tubes which were then treated as follows :-— Tubes 1 and 2, added to bouillon one drop of fresh Crotalus venom; mouth of tubes plugged with cotton. ‘Tubes 3 and 4, prepared same as last, but tube left open (no cotton plug). ‘Fubes 5 and 6, added one grain of Crotalus peptone. ‘Tube closed by cotton plug. Tubes 7 and 8, same as last, but tubes left open. Tubes 9 and 10, added one grain of Crotalus globulin. Tubes closed. Tubes 11 and 12, same as last. Tubes open. Tubes 13 and 14, a pure bouillon, nothing added to it. Tubes closed. Tubes 15 and 16, same as last, ‘Tubes open. ‘Twenty-four hours later the bouillon in all the test tubes which originally was perfectly clear had become cloudy except tubes 13 and 14 (which contained thie sterilized pure bouillon plugged well with cotton). On the third day of the experiment tubes 3 and 4 (fresh venom, tubes open) showed well-pronounced putrefaction of the bouillon. Slight putrefactive changes were subsequently observed in the remaining tubes (except 13 and 14) in the following order :— On the fourth day, tubes 7 and 8. On the fifth day, tubes 11 and 12, also in tubes 15 and 16. On the seventh day all the plugged specimens were examined, and all showed PATHOLOGY. 135 more or less putrescence except the tubes with the pure bouillon as stated. Of these closed test tubes, however, tubes 1 and 2 (the fresh venom) showed the putrefactive changes to be much more pronounced than in the remaining tubes ; but as we have seen putrefaction ensued much sooner in the tubes that were open (tubes 3 and 4). As all the tubes showed putrefaction more or less, it is presumable that the peptone and globulin accidentally contained bacteria, these substances not having been sterilized. at the commencement of the experiment. The contents of the tubes examined microscopically during and at the end of the experiment showed the presence of bacteria of putrefaction in direct proportion to the putrefaction changes. Imperfect as this experiment may be, it appears to establish the fact that fresh venom promotes putrefactive changes comparatively more rapidly than the venom peptone and globulin, but it also shows further that this power to produce putres- cence is very much aided by the action of the air, and depends upon the presence of bacteria contained in that air or in the venom. It was also evident that putre- faction was considerably retarded in all the tubes that were plugged by the cotton, and further that unplugged tubes containing sterilized soup, and exposed to contami- nation from air showed also putrefaction but at a later date. Putrefaction Experiment with Muscular Tissue and Venom.—The following rough experiment also appears to show that putrefactive changes develop in dead animal tissues much more rapidly in the presence of venom than without it. Experiment.—A few drops of a solution of dry Crotalus venom were poured upon a small piece of fresh muscle just removed from the thigh of a rabbit and placed in a covered glass beaker. A similar preparation but without the addition of venom was made in a second covered beaker. Temp. 70° to 80° F. Putrefactive changes began to appear in the specimen treated by the venom after twenty-four hours, and after seventy-two hours were quite far advanced. Under the microscope the muscular tissue showed necrotic alterations very similar to those (to be described later) as occurring in experiments upon the living muscle. A multitude of dumb-bell-shaped rod bacteria, some large bacilli and the micrococci of the venom enormously multiplied, were seen in the decaying muscular substance. In the specimen of muscle not treated by venom, putrefactive changes were delayed to the fifth day and then appeared to be much less conspicuous, showing but few bacteria. ‘The muscle fibres were uniformly cloudy and degenerated but not broken down in the peculiar manner caused by venom. Experiments with Bouillon and Venom in Sealed Glass Bulbs, Venom being thoroughly Sterilized.—More satisfactory and conclusive results were obtained from the following experiments :— A number of small glass bulbs were filled with sterilized bouillon after the well- known method of Dr. Sternberg, and after being thoroughly resterilized by boiling the following preparations were made :— To each of six bulbs was added one grain of dry Crotalus venom, the venom having been previously subjected to sterilization in a dry heat at 110° C. for one 136 THE VENOMS OF CERTAIN THANATOPHIDES. hour. The bulbs were then hermetically sealed by melted glass. The bouillon in these tubes (with sterilized venom) remained perfectly clear and free from bacteria. Microscopical examination was made at various periods, the last time after eighteen months when it was still perfectly clear and showed no signs of putrefaction. A similar result was obtained in an experiment with another set of six glass bulbs filled with bouillon, and to which some Moccasin peptone, previously sterilized, was added. ‘These bulbs looked somewhat cloudy, but on examination of the con- tents eighteen months later no bacteria, and no putrefactive changes were noted. As a control experiment six bulbs filled with pure sterilized bouillon were kept for a similarly long period, and they all remained clear and free from change; while a few bulbs filled with unsterilized bouillon showed great cloudiness, bacteria, and putrefactive change. Ath. Culture Experiments.—The study of the morphology of the bacteria inhabit- ing the venom was next undertaken. To this end numerous culture experiments to isolate the bacteria from the venom were made. As stated before, the perfectly fresh venom contained only one form of these vegetable organisms, the micrococci, and only to these latter attention was paid; the rod bacteria and bacilli not appearing except in venom which had began to putrefy. The micrococci contained in the venom showed the following behavior in pure ~ cultures: Of culture soils, the peptonized gelatine prepared after the formula of Koch proved to be quite suitable. The isolation of the micrococci was made after the methods of Sternberg and of Koch, as adopted in the pathological laboratory of the University of Pemsylvania. For gelatine culture a minute quantity of venom was smeared on the surface of the solidified jelly contained in a sterilized, small, flat, well covered glass vessel. The micrococci liquefied the jelly, an effect not peculiar to all bacteria. After twenty-four hours all over the inoculated surface of the jelly were seen small turbid drops which contained the micrococci. With a sterilized platinum wire the micrococci from one of the liquefying specks upon the first culture were transplanted to the jelly in a second culture vessel. From this second generation a minute quantity was transplanted to a third and fourth culture vessel. The fourth and all the later generations yielded usually a pure crop of micrococci. : In impure cultures dumb-bell-shaped bacteria and sometimes large bacilli were met with. ‘These, however, could not be said to be peculiar to venom, as they are never found in fresh venom. It may, therefore, be concluded that these cultures represent the micrococci peculiar to, or at least those constantly inhabiting venom. More- over, the micrococci in these cultures whenever they were successful, were the only bacterium seen and were fully identical as to shape, measurement, and be- havior to aniline dyes with those found in the fresh venom. Much better crops of the venom-micrococci were obtained in bouillon cultures in Sternberg’s glass-bulbs. The micrococci grow more rapidly and better in these bulbs, because the bouillon can be heated up to the more suitable temperature of 40° C.; while the jelly cultures could not be warmed to such a degree without melt- ing solid gelatine. A variety of other culture soils and methods of isolation were PATHOLOGY. 137 employed in these experiments, but their description here is unnecessary as not being sufficiently related to the points at issue, In relation to the morphology of the micrococci it may be added, that they measure on the average 7545, of an inch in diameter; they often appear in pairs, but most commonly in zoogleea masses. ‘They show a distinct aureole, such as is met with in various forms of micrococci.’ These aureoles have lately been erroneously described by Friedlander, as peculiar to certain “ specific” micrococci in croupous pneumonia. In conclusion, it might be said that the venom micrococci do not appear to differ from the micrococci found in the saliva of men and other animals. In order to test whether the venom-micrococci were in any way specific or patho- genetic, and whether they form, or contribute to, the virulence of the venom, inocu- lations with pure cultures of the micrococci were made upon animals. As these experiments gave entirely negative results, it is superfluous to enter into details. Suffice it to say that large quantities of the pure micrococci from a sixth generation were injected, in various manners, into rabbits, cats, pigeons, and white rats, but without fatal results; or without producing any other lesion than occasionally local abscesses, or later on, metastatic abscesses. Sometimes the so- called “miliary tuberculosis of animals” was produced by inoculating with the venom-micrococci. No signs of any lesions resembling those of venom poisoning were observed. Experiments made to Study the Anatomical Changes produced by the Venom in liv- ing Animals. Naked Eye Appearances.—Very many years ago Dr. Weir Mitchell described two forms of venom poisoning—rapid or acute, and slow or chronic. ‘To the latter appear to be relegated by him all those cases in which death is protracted beyond a few hours. This convenient division is justified by certain differences in the mode of termination of venom poisoning, and by the macroscopic and micro- scopic appearances of the lesions induced. In the most rapid poisoning, there is frequently nothing appreciable to the naked eye beyond the slight local lesion or here and there minute capillary hemorrhages, when death has been delayed beyond a minute. In examples of chronic poisoning both the local and the systemic changes are enormously more extensive. When animals were subjected to chronic poisoning they were kept under the influence of narcotics, since it had been learned that these agents did not affect the results. No Cobra venom was employed in this series, but only the pure or dried venoms of our own serpents, or else some one or other of the constituents of these poisons. The following tables relate the experiments made, and the more striking mor- phological changes :— * See “Memoir on Diphtheria,” Report to the National Board of Health, 1882, by H. C. Wood and H. F. Formad. 18 June, 1886, 138 THE VENOMS OF CERTAIN THANATOPHIDEA. RAPID POISONING. Liffects of Venom when Injected Hypodermatically into or Applied otherwise to the Tissues of the Living Animal. No. of | Animal | Form and quantity | Time of Local lesion. Condi- | Changes in thorax, abdomen, REMARKS. expt. | used. of yenom, and death. tion of brain, and membranes. where introduced. blood. 1 | Pigeon | Crotalus venom, Killed | Moderately sized,| Coagu- | All internal organs congested; | This animal was killed fresh, 7 grain in- | after 5 | dark hemorrha-| lable no other changes visible; no} before the full effects of jected into pecto- |minutes| gic swelling and red | ecchymoses perceptible the venom. ral muscle 2 | Pigeon | Same as last Died | Very dark colored} Less co-| Organs only moderately con- | For the details and the his- in 15 hemorrhagic agulable| gested, but there were nu-| tological appearances, see minutes] swelling and merous small subpleural,| the next chapter. The quite subperitoneal, and slight | studies of the changes dark subpericardial ecchymoses in muscular tissue were mostly made from this experiment. 3 | Pigeon | Moccasin venom, Died | Profuse hemor- Liquid | Ecchymoses in nearly all or- fresh, 1 drop in- | after 1 rhage all over very gans, quite marked in arach- jected into peri- hour peritoneal dark noid and at base of brain ; toneum and 50 | cavity some so small as to be visi- minutes ble only by microscope. Ex- treme congestion 4 | Pigeon | Crotalus venom, Died | Same as last Liquid | Hemorrhages only subperito- fresh, 1 drop into} in 25 dark neal, other organs merely peritoneum minutes congested 5 | Rabbit | Moccasin venom, 9 Subperitoneal Coagu- | No changes beyond local injected into peri- | minutes} hemorrhages lable on | lesion toneum exposure 6 | Pigeon | Peptone, injected 35 Hemorrhagic Liquid | No visible changes, except all| Changes in muscular tis- into pectoral minutes} swelling organs congested sue similar to those pro- muscle duced by fresh venom, but far less blood effused. 7 | Pigeon | Moccasin venom, 5 Hemorrhages in | Slightly | Membranes of brain and brain fresh, injected minutes} arachnoid and | coagu- substance peripherally soak- into cavity of brain tissue lable ed with blood; other organs skull congested 8 | Rabbit | Moccasin venom, 1 Lung infarcted | Same as|No changes, except in lung | See specimen and deserip- fresh, } grain in- | minute} by blood last and some subpericardial ec-| tion in chapter on histo- jected into lung chymoses logical changes. 9 | Rabbit | Peptone, $ grain 43 Ecchymosis Liquid | Other organs not visibly into liver and minutes| locally only dark affected peritoneum 10 | Rabbit | Same as last Killed at| Slight subperito-| Dark, | No systemic changes. the end | neal ecchymoses} but co- of 1 hour agulable 11 | Pigeon | Peptone, 1 grain 20 Local ecchymoses| Ditto } ( Histological changes into peritoneum | minutes] slight | No notable changes in other } similar to those pro- 12 | Pigeon | Globulin, } grain The same as last} Ditto |{ organs duced by fresh into peritoneum | minutes { venom. 13 | Cat Crotalus globulin, 40 Profuse ecchy- Slightly | Hemorrhage only local $ grain into peri- | minutes| mosis coagu- toneum lable 14 | Cat Crotalus peptone, | 1 hour | Same as last, but} Liquid | Hemorrhage only local, also | | Microscopic examination 3 grain into peri- | and 20 | less marked extreme congestion of all|‘+ made of every organ. toneum minutes organs The details will be 15 | Rabbit | Crotalus globulin, Killed | Same as last Red and| Nothing peculiar beyond the given hereafter. 3 grain into peri- | in 10 coagu- local lesion toneum minutes lable J 16 | Rabbit | Dry Crotalus Killed | Same as last Same as | Same as last venom, $ grain in| in 10 last watery solution, | minutes into peritoneum 17 | Rabbit | Dry Crotalus) Died | Same as last Very | Ecchymoses in all organs ex-|Some of the ecchymoses venom, $ grain in| 1 hour dark, amined. Profuse ecchymo-| were so small as to be watery solution, | and 25 liquid ses in peritoneum, also snb-| visible only on micro- into peritoneum | minutes pleural, subarachnoid, and} scopical examination. subpericardial. Liver, which was injured by the syringe, showed a large hemorrhagic infarction PATHOLOGY. 139 RAPID POISONING.—Continuep. No. of | Animal | Form and quantity | Time of Local lesion. Condi- Changes in thorax, abdomen, REMARKS. expt. used. of venom, and death. tion of brain, and membranes. where introduced. blood. 18 | Cat Dry Crotalus Died | Same as last Same as| Peritoneal cavity contains a| Cats appear to resist the venom, 1 grain in | 5} hours last good deal of liquid blood;} effects of venom much watery solution, hemorrhage at base of brain] longer than the other into peritoneal (subarachnoid); no other] animals used in this re- cavity lesion noted; organs rather| search. anemic. Heart empty, con- tracted 19 | Cat Peritoneumopened,}| Died | Hemorrhagic in- | Partly | Peritoneal hemorrhage; or-| It appears that when the (chlor- | mesentery exposed| after 4 | filtration, quite | coagu- gans anemic mesentery is exposed and alized) | uninjured, in hours extensive, but lable not injured the animal moist chamber, and 35 | came on very survives much larger ap- and smeared re- |minutes| slowly plications of venom than peatedly with a if venom be injected into solution of dry an unopened peritoneal venom, using not cavity. Very small quan- less than 5 grains tities of venom appear to of venom kill in the latter case. For further experiments of this character, Mechanism of rhages. None of the cases in the table exhibit instances of the greatest possible rapidity of death. Dr. Mitchell has seen a pigeon die within ten seconds from a hypodermatic injection of pure Crotalus venom, In such a case there is positively no lesion, and the blood is solidly coagulated. In most cases very soon after injection of the venom in either of its forms, the time varying from a few minutes fo a few hours, according to the kind of animal and the quantity of venom used, there appears a swelling at the point of injection with intense violet-black discoloration of the skin, which gradually extends over an area of several square inches. On making an incision into the tissues in the immediate neighborhood of the injection, they are found to be soaked with extravasated blood. ‘This is often all that is visible if death has occurred soon; but if it has been postponed for a short time, then in tissues distant from the place of the injection, extravasations to a smaller extent were often found. Most pro- nounced and most frequent are the ecchymoses below serous membranes (subpleural, subperitoneal, and subpericardial) ; in fact the whole organism is deeply affected, the tissues being congested and presenting a much darker appearance than normal. The blood does not seem to coagulate readily within cavities or interstices of the body unless death follows almost instantaneously. In cases which live longer, the blood remains commonly in a liquid state, or coagulates imperfectly, and then only after being exposed to the air, resembling in this particular the state of that fluid observed in conditions of asphyxia. REMARKS, 140 THE VENOMS OF CERTAIN THANATOPHIDES. SLOW POISONING. Effects of Venom upon the Tissues of the Living Animal. No. of | Animal | Form and quantity | Time of Local lesion. Condi- Changes in internal organs. expt. used. of venom, and death. ; tion of where introduced. blood. 20 | Pigeon | Copper globulin, 13 Large, dark gan-| Liquid | Subpericardial ecchymoses and 2c.c. (equal tol} hours | grenelikeswell-| and pericardial effusion. Red tinged gram fresh venom) ing of chest ; dark serum in peritoneal cavity. Heart injected into pec- muscle disinte- empty. Lungs and pleura full toral muscle grated of ecchymoses. All the organs congested 21 | Rabbit | Unknown, but very | 9 days | Dark gangren- Liquid | Numerous minute hemorrhages be- minute quantity ous swelling and low serous membranes, seen also of Crotalus venom dark at base of brain in right posterior injected in back fossa. The organs rather anemic and softened ; 22 | White | Crotalus venom, 2 days | Hemorrhagic Slightly | Organs congested, softened ; noth- rat dry, $ grain in- and 7 peritonitis coagu- ing else peculiar found; small, jected into abdo- hours lable, loose, red clot in right side of men dark heart 23 =| Cat Crotalus venom, 9 days | Skin slough over | Liquid | All internal organs softened and dry, 1 grain in- and 2 local lesion, dark, ill) highly ecchymosed and congested. jected into right hours which is dark, | smelling] Feces and urine bloody. Hemor- thigh hemorrhagic, rhage at base of brain, and min- and gangrenous ute blood specks in pericardium. Heart quite atrophied and softened 24 | Pigeon | Quantity unknown,| 14 days| Atrophy, with Liquid | Hemorrhages indicated by deposits injected into pec- pigmentation of} and of blood pigment in the tissues. toral muscle the pectoral very All the organs in a state of atro- muscle injected | dark phy and softened, resembling acute yellow atrophy in man. Serous sacks all distended with bloody serum. Heart empty, and although contracted quite soft All these autopsies were made immedi- ately or quite shortly after death. For changes in blood, see details in text. For further details of the histological changes, see text. N. B.—Gangrenous changes in the local lesion are usually more pronounced in the ‘‘Slow”’ than in the Rapid form of venom poisoning. The following lesions may be mentioned as peculiar to retarded or slow poison- ing: Rigor mortis often absent. does not readily acquire the scarlet-red color when exposed to the air, are prominent blood-stained effusions in all the serous sacks. and feeces often bloody. ous than in the rapid’ poisoning. The blood, usually diffluent, is very dark and There (Plate V.) Urine Hemorrhages beyond the local lesion much more conspicu- The remote lesions of slow poisoning resemble very much (morphologically) the primary local lesion, but are not so extensive or so well defined. of acute septic poisoning. the manifestations of rapid and slow poisoning, nevertheless the division is in prac- tice convenient. One case of very protracted slow poisoning was observed in a pigeon which had been injected with venom in the pectoral muscle. Slow Poisoning.) Instead of the usual gangrenous change there was seen in this case after the lapse of two weeks a decided dry atrophy of the muscular tissue about the wound. Its fibres were greatly diminished in size as compared with the opposite unaffected In general the conditions of slow venom poisoning resemble those It is very often impossible to draw a distinct line between (See Experiment 24, ‘Table PATHOLOGY. 141 muscle, and many of them were entirely disintegrated, as was evident from the remnants of the muscular fibres and the granular material which took their place between the interstices of the connective tissue. ‘This granular material was seen throughout the specimen, some of it being of a brown tint, and probably repre- senting disintegrated blood corpuscles. ‘The internal organs were all in a state of atrophy, more particularly so the liver, the tissues of which under the microscope bore a striking resemblance to acute yellow atrophy. The serous sacks were all largely distended by blood stained serum. ‘The heart muscle was also in a condi- tion of atrophy, its chambers empty, and the blood dark and not coagulable. Blood examined microscopically showed appearances to be mentioned shortly. The Effects of certain Venoms on the Coagulability of the Blood.—One of the most interesting differences in the action of the venoms of the Rattlesnake and Cobra and which was pointed out some years ago by more than one observer, is that the former venom partially or completely destroys the coagulability of the blood, while the venom of the Cobra has no such marked effect. The blood of animals poisoned with Crotalus venom is usually thin and dark, the clots form slowly, and are very soft and easily broken up. Some direct studies were made to test more accurately this interesting property of the Crotalus venom, and it was thus observed that it is not peculiar to the poison of this genus, but is also a characteristic of the Moccasin. 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Zur Behandlung des Schlangenbisses durch subcutane Injection von Liquor ammon. caust. Berl. klin. Wehnsehr., 1873, X, 330. Bite from the hooded snake—cobra de Capello—rapid death; autopsy. Lancet, Lond., 1852, II, 397-400. PRIov. Romz QuUAIN. DE QUATREFAGES. Note sur le permanganate de potasse, considéré comme antidote du venin des serpents, 4 propos d’une publication de M. J. B. de Lacerda. Compt. rend. Acad. d. se., Par., 1882, XCIV, 488-490. Also: France méd., Paris, 1882, I, 319- 321. Raz (W.). Ona case of snake bite treated by ammoniacal injection. Austral. M. J., Melbourne, 1869, XIV, 242-244. Rawpw (T.8.). Observations on the action of snake poison on the blood. Austral. M. J., Melbourne, 1867, XII, 353-361. Bey bali OlGaPAsbareves RANKEN (J.). An account of some experiments made upon dogs bitten by the cobra de Capello, or coluber naji. Edinb. M. and 8. J., 1822, XVIII, 231-239, Vergiftung eines Knaben durch den Biss der Vipera berus. Ztschr. f. Chir. vy. Chir., Osterode, 1841, I, pt. 2, 36-39. Recovery from the bite of the cobra de Capello. Lancet, Lond., 1859, II, 13. Rep (F.). Lettera sopra aleune opposizioni fatte alle sue osservazioni intorno alle vipere. Scritta alli signori Allessandro Moro e Abate Bourdelot. 31 pp. 4°. Firenze, 1685. Repi (F.) Osservazioni intorno alle vipere. Eda lui scritte in una lettera all’ illustris- simo signor Lorenzo Magalatti. 1 p.1., 91 pp. 4°. Firenze, 1664. REIMONENQ. Un jeune garcon de dix-neuf ans qui avait été mordu par un serpent. J. de méd. de Bordeaux, 1853, 748-750. Remede contre la morsure de vipere. [Mém. Acad. ry. d. se. 1747.] Collect. acad. d. mém., etc , Par., 1785, X, 445. ReEmeEDIOS Monrerro (J.). Do permanganato de potassa contra o veneno das cobras. Gaz. med. da Bahia, 1881-2, 2. S., VI, 197-199. Remer (W.). Folgen des Bisses einer giftiger Schlange. J. d. prakt. Heilk., Berl., 1814, XXX VIII, 1 St., 47-60. Renecer (J. R.). Ueber die Wirkung des Bisses der siidamerikanischen Giftschlan- gen, und die von mir dagegen angewandte Heilmethode. Arch. f. Anat. u. Physiol., Leipz., 1829, 271-298. Also, trans.: J. compl. du dict. d. sc. méd., Par., 1830, XXXVII, 246-264. Report of the Commission appointed to investi- gate the influence of artificial respiration, intravenous injection of ammonia, ete., in Indian and Australian snake poisoning. Indian Ann. M. Sc, Calcutta, 1875, XVII, 191-252, 4 pl.; with Appendix, Nos. 1-3, pp. I-XCVIII. Report of the special committee on the subject of snake poisoning. Austral. M. J., Mel- bourne, 1877, X XI, 104, 151, 184. RATH. Return showing the number of deaths from snake bites, in the year 1869, in the Province of Bengal. Population, including Orissa and BIBLIOGRAPHY. Assam, 48,358,134. Indian M. Gaz., Cal- cutta, 1870, V, Suppl., pp. 1-4. RicHarps (Y.). Experiments on snake poison. Indian Ann. M. Se., Calcutta, 1870, No. XXVIII, 177-202. Ricwarps (V.). Case of cobra bite. M. Gaz., Calcutta, 1871, VI, 130. RICHARDS (V.). by artificial respiration. Caleutta, 1872, VII, 247. Ricwarps (V.). Snake poisoning antidotes. The nature of snake poison, and its action on the blood. Indian Ann. M. Sc., Calcutta, 1872-3, XV, 163-176. Ricwarps (V.). Experiments with reputed anti- dotes and on artificial respiration in snake poisoning. Rep: on . . . san. improve. in India, 1872-8, Lond., 1878, 264-275. Ricwarps (V.). Dr. Fayrer’s treatment of snake bite by artificial respiration. Indian M. Gaz., Calcutta, 1873, VIII, 118-120. Ricuarps (V.). The treatment of snake bite by intravenous injections of ammonia. Med. Times and Gaz., Lond., 1873, I, 639. RicHarpDs (V.). Experiments with strychnine as an antidote to snake poison. Med. Times and Gaz., Lond., 1874, I, 595-597. Ricwarps (V.). Experiments with snake poison. Indian Ann. M. Sc., Caleutta, 1873-4, XVI, 285-301. RicHarps (V.). Report on the snake bite cases which occurred in Bengal, Behar, Orissa, Assam, Cachar, ete., during the year 1873-4. Indian M. Gaz., Calcutta, 1876, XI, 96-100. Ricuarps (V.). A fatal case of snake bite, in- travenous injection of ammonia; remarks on the application of the ligature in snake bite. Indian M. Gaz., Calcutta, 1876, XI, 320. Ricwarps (V.). A case of snake poisoning; no treatment; recovery. Lancet, Lond., 1878, I, 530. RicHARDs (V.). Snake poisoning and its treat- ment. Indian M. Gaz., Calcutta, 1880, XV, 309. Ricwarps (V.). A case of snake bite; hypoder- mic injection of permanganate of potash. Indian M. Gaz., Calcutta, 1882, XVII, 44. Ricuarps (V.). Further experiments with per- manganate of potash, liq. potasse, and Indian Treatment of snake poisoning Indian M. Gaz., 175 iodine in cobra poisoning. Indian M. Gaz., Calcutta, 1882, XVII, 199-202. Ricuarps (V.). Notes on Dr. Wall’s monograph on cobra and Daboia poisons. Indian M. Gaz., Calcutta, 1882, XVII, 239, 259. Ricuarps (V.). Permanganate of potash and liquor potassz in snake poisoning. Lan- cet, Lond., 1882, I, 1097. Ricwarps (V.). Dr. Badaloni on the perman- ganate of potash. Lancet, Lond., 1883, II, 461. RicHArpson (J. F.). Rattlesnake poison. Phila. M. Times, 1879, IX, 306. Ricutrer (M.). Vergiftung eines Midchens durch den Biss der Coluber berus. Ztschr. d. deutsch. Chir.-Ver. f. Med:, chir. u. Geburtsh., Magdeb., 1850, IV, 9-15. Rivers (G. M.). The rattlesnake, its poison and antidote. South. M. Ree., Atlanta, 1874, LV, 505-513. Rosert. Morsure d’une vipére. Bull gén. de thérap., etc., Par., 1834, VII, 307-309. Rogson (T.). Case of snake bite, in which bleeding was used as an auxiliary. Tr. M. and Phys. Soc., Caleutta, 1835, VII, 2. pt., 480-482. Rogers (8.). Case of snake bite. M. J., 1839, I, 231. Rosrt (W.G.). On snake bites. India J. M. and Phys, Se., Calcutta, 1836, N. S., I, 448. Rousseau (Ll. F. E.). Expériences faites avec le venin d’un serpent & sonnettes ( Crotalus horridus.) J. hebd. de méd., Par., 1828, J, 291-296. Roy (G. C.). Experiments with cobra poison. Indian M. Gaz., Calcutta, 1876, XI, 313. Roy (G. C.). Remarks on the action of snake poison on the blood. Indian M. Gaz., Calcutta, 1877, XII, 315-317. Madras Q. Roy (G. C.). nes. Anagenesis, or the Genesis of Progressive Characteristics, 71-74. Catagenesis, or the Genesis of Retrogressive Characteristics, 74-80. Differential Characteristics, 80-84. © - VAR CHOROGICAT PAN DS AUNAD IED ATTONS eihee Gl) Sele) =) fee Geo es ke . 85-119 Remarks, 85-89. Psiloceras and Caloceras, 89-93. Weehneroceras and Schlotheinia, 93-95. Vermiceras, 95, 96. Arnioceras, 96, 97. Coroniceras, 97-99. Agassiceras, 99,100. Asteroceras, 100, 101. Oxynoticeras, 101-103. Fauna of South Germany, Table I., 103. Fauna of the Cote d’Or, Table II., 103, 104. Fauna of the Rhone Basin, Table III., 104-106. Fauna of England, Table IV., 106. Fauna of the Proy- ince of Central Europe, Table V., 106-108. Fauna of the Province of the Mediter- ranean, Table VI., 108-112. Summary, 112-119. V. DEscRIPTIONS OF GENERA AND Species OF ARIETIDH ...... . =. 120-291 Radical Stock, 120-125. First, or Psiloceran Branch, 120-125. Psiloceras, 120- 124. ‘Tmegoceras, 125. Plicatus Stock, 125-161. Second, or Sehlotheimian Branch. 125-136. Wehneroceras, 125-127. Schlotheimia, 127, 128. Third, or Vermiceran Branch, 136-161. Caloceras, 136-154. Vermiceras, 154-161. Levis Stock, 161-221. Fourth, or Coroniceran Branch, 161-194. Arnioceras, 161-174. Coroniceras, 174-194. Fifth, or Agassiceran Branch, 194-214. Agassiceras, 194-200. Asteroceras, 200-214. Sixth, or Oxynoticeran Branch, 214-221. Oxynoticeras, 214-221. Na ne eee ee mee SS el Le 8 oR % my ORTHOCERAS ELEGANS, APEX AND PROTOCONCH . OrtHocerAs potirum, APEX AND PRovToconcH. . ORTHOCERAS TRUNCATUM, TruncateD Exp ORTHOCERAS UNGINS, CICATRTX AND APEX OrTHOCERAS POLITUM, APEX .... . WaHNEROcERAS EMMRIcHT. ... . CALocERAs PAGE 17 AGN (AUST (50 a CanocreRAS NEWBERRYI ..... . AND PLUG (CATO CHRIS OlR 1 O:Ni Ie ARNIOCERAS NEVADANUM .. .. . .- ASTEROCERAS OBTUSUM, VAR. QUADRAGONATUM LIST OE CUMS SING BER Ob ere yaar ia tee 2 REL) ae lec er UG Bise. A “ce 6-8, (G5 te Sale #1819, « je C2022 de 12’ . 23, 24, a ‘ bi OG re mca PREFACE. pe is a common mistake to designate my classification as “embryological.” It will be found by those who read these pages, that the whole life of the individual, and all its metamorphoses, have been deemed essential standards for the estimation of affinities. Even the degradational meta- morphoses of old age are used as characteristics of value in the generic descriptions; it is properly speaking an ontological classification. The researches were conducted almost wholly in Museums, because it was found impracticable to study stratigraphical superposition in the field. This part of the work has already been accurately done by local geolo- gists, and my notes were largely made upon their collections. More extended studies might have made the work more accurate than it is, but this was not possible for me. I desire to record my deep sense of obligation to the late Prof. Louis Agassiz, under whose direction my studies upon the Arietidae were begun. ‘His instruction and advice were none the less valuable because we differed in theoretical views; to him I owe the methods of observation which are used in all my work. His son, Alexander Agassiz, has also laid scientific men in this country under heavy obligations, and this essay could not have been completed or published but for his sympathy, and for the liberal manner in which he has sustained by large personal sacrifices the collections and the cause of scientific research in the Museum of Comparative Zodlocy. Professor Langley, Secretary of the Smithsonian Institution, has shown the greatest consideration and courtesy, and in undertaking the speedy pub- lication of this memoir after the Museum of Comparative Zovlogy had been obliged for want of funds to postpone its issue indefinitely, has saved the results from becoming antiquated before they were made public. My principal studies outside of this Museum were made in the Museum of Stuttgardt, and there I received unwearied attention and help from Prof. Oscar Fraas, and the use of superb collections. Professor Quenstedt of Tiibingen gave me the benefit of much valuable information, and threw open his collections without reserve, and I am indebted for similar favors to Prof. Guido Sandberger at Wiirtzburg, Prof. Karl Zittel of the Museum at Munich, and to Professor Mésch at Ziirich. The late M. Barrande, Professor Gaudry and his assistant Dr. Fischer of the Jardin des Plantes, ( vii ) vill PREFACE. Professor Hébert and his assistant M. Munier-Chalmas of the Sorbonne, Paris, were equally kind and liberal. I desire also to thank M. Collenot, M. Bréon, and Dr. Bochard, for their kind attention and the free use of the collections at Semur. Professor Owen and Dr. Henry Woodward of the British Museum, Mr. Etheridge of the Geological Museum, the authori- ties of the Bristol Museum, and Dr. Thomas Wright, gave me_ similar opportunities for study, and Mr. Marder at Lyme Regis assisted me in the field. Prof. Jules Marcou has materially aided the work by the loan of rare books not obtainable elsewhere, and I am also indebted to Prof. J. D. Whitney for similar loans from his library. Professor Emerson of Amherst has given me valuable information, and the use of his collection. I was unfortunate in finding the curators of collections either absent or sick at Hanover and Heidelberg; but in all practicable cases ample opportunities for study were given me, except at the Museum of York, England, where unyielding regulations prevented access to the interior of the cases, and my identifications there were consequently made without handling the speci- mens. I am also indebted to Professor Cope and Dr. John A. Ryder for the results of investigations which have thrown much light upon vexatious questions of theory, and which have not been properly repre- sented by quotations in the text of this work, the general remarks having been necessarily cut down to the narrowest possible limits. - The essay on “ Fossil Cephaiopods in the Museum of Comparative Zodlogy ” was written in large part as an introduction to this monograph, but for obvious reasons has not been used. The following conclusions, copied with some emendations and corrections from that essay, may be useful, however, in giv- ing the reader a view of the theoretical opinions entertained by the author. 1 1. Law of Morphogenesis.— We have endeavored to demonstrate that a natural classification may be made by means of a system of analysis in which the individual is the unit of comparison, because its life in all its phases, morphological and physio- logical, healthy or pathological, embryo, larva, adolescent, adult, and old (ontogeny), correlates with the morphological and physiological history of the group to which it belongs (phylogeny ). 2. Organic Equivalence.— All new characteristics, even those which are purely mechanical reactions of the tissues, arise in a similar manner, as reactions due to the exciting agency of the more general or more localized physical causes. They are there- fore necessarily, and because of this mode of origin, the corresponding organic, or suitable complementary equivalents of these physical causes, both structurally and functionally. 3. After their origin, however, and during their subsequent history, organic equiva- lents or characteristics are divisible into two categories: those which become morpho- logical equivalents, and are essentially similar in distinct series, and those which are essentially different in distinct series, and may be classed as morphological differentials. 4. Morphological Equivalence.—In the different genetic series of a type derived from one ancestral stock there is a perpetual recurrence of similar forms in similar succes- sion, which are usually called representative and often falsely classified together, though they really belong to divergent, genetic series. 1 Proc. Am. Ags. Ady. Sci., XXXII, 1883. PREFACE. ix 5. These forms and their similar characteristics are not derived by direct inheritance from the common ancestor, in which all the forms are necessarily similar and primitive, but originate everywhere independently of hereditary influences in the different series, and also in all formations independently of chronological or chorological distribution. 6. This evolution of similar morphological changes in the forms of different genetic Series must be regarded as the similar reactions or efforts of a common organism in direct response to similar generally distributed physical causes active in the same habitat, and are therefore necessarily similar to each other, though in different genetic series. As a whole, they may be said to express the general tendencies of modification, due to the efforts of the common-radical and common organization while spreading in all direec- tions and in different genetic lines to respond to similar physical causes, and meet their requirements with suitable changes. They are, therefore, structural equivalents of each other in different series, and functional equivalents of the general requirements of the environment or habitat, or, in other words, purely physical selections. 1. Morphological Difference. — Differentials are absent in the first members of series, on first appearance in their descendants transient, but afterwards tend to become inva- riable, or fixed in the stock or series, being perpetuated by direct inheritance in succes- Sive generation’, species, ete. They finally often disappear in the retrogressive or highest and last occurring members ofeach series, or in aberrant forms when on the same level. 8. They have no determinate mode of succession, but are usually more or less isolated modifications, und arise first in individuals or varieties, but afterwards become characteristic of species, and finally of the major part of the direct line in species, or descendent series. 9. They are, therefore, strictly adaptive, variable characteristics, and not directed in their occurrence or development by any more or less invariable law of successive modi- fication, as are the morphological equivalents. We have failed in finding any differentials of great importance whose prepotence as hereditary characteristics could not be accounted for by the law of use and disuse in connection with habits. The differentials of small series, species, genera, and families, which we have not been able to analyze thoroughly, may be due to the action and reaction of individual animals upon each other, or, in other words, to natural selection. 10. Differentials, therefore, can be separated from-other characteristics of the same parts by careful observation and close analysis of their behavior in series, but cannot be specifically predicted from the study of other series; whereas, morphological equiva- lents can be predicted with the same certainty as the recurrence of cycles in physical phenomena. Thus we can say of any new series of Nautiloids or Ammonoids, that, the habitat remaining similar, they will, whenever or wherever found, tend to develop arcuate, coiled, close-coiled, or discoidal and finally involute forms in progressive series, and reverse this process in retrogressive series. 11. Acceleration in Development.— All modifications and variations in progressive series tend to appear first in the adolescent or adult stages of growth, and then to be inherited im successive descendants at earlier and earlier stages according to the law of acceleration, until they either become embryonic, or are crowded out of the organization, and replaced in the development by characteristics of later origin. 12. Geratoloyy.— Modifications which tend to appear in the old age of the individual of progressive series correlate with the modifications taking place in pathological series of all grades, and in geratologous and retrogressive forms of all kinds, however progres- sive they may be in certain characteristics. Geratologous forms, therefore, show that the development of retrogressive characters has been stimulated so as to take the place of the hereditary progressive, thus either partially or completely replacing them. Partial replacement is often accompanied by the early development of hereditary progressive characteristics. x PREFACE. 13. Acceleration in Degeneration. — Geratologous forms may, therefore, be the highest members of progressive series, or the terminal members of retrogressive series, and the stimulation of the development appears to take effect upon both progressive and retro- gressive characteristics ; thus producing, at the same time and in the same animal, first, the earlier development of some of the progressive characteristics combined with geratologous characteristics ; secondly, the earlier development of geratologous characteristics and their fusion with larval characteristics, which occasions the complete replacement of progressive characters, and occurs only in the extreme forms of retrogressive series, and in parasites. 14. The law of acceleration in development seems,’therefore, to express an inya- riable mode of action of heredity, in the earlier reproduction of hereditary characteris- ties of all kinds, and under all conditions. In progressive series it acts upon healthy characteristics, and appears to be an adaptation to favorable surroundings, and in retro- gressive series upon pathological characteristics, and is probably an adaptation to un- favorable surroundings, usually leading to the extinction of the series or type. 15. The Three Phases of Development. —In following up series, it has been found that the development of ancestral forms is simple and direct (Epacme); that of their more specialized descendants becomes gradually indirect (Acme), acquiring complicated inter- mediate or larval stages; and that of the terminal retrogressive or geratologous and pathological forms becomes again more or less direct (Paracme. ) 16. The introduction of adaptive larval stages into the history of individual develop- ment in any series appears to be due to the direct exciting action of the surroundings, and their absence or subsequent suppression to some physical agency, changes of habit, or protection, or pathological causes. All of these causes must, however, be considered as similar in their effect upon the young. They are stimulants, producing acceleration or excessively rapid development of the ancestral progressive characteristics, or of the retrogressive, or primitive larval characteristics inherited from the progressive forms. 17. This agreement in the mode of development of the individual according to its position in the history of the group completes the correlations which exist between the history of the individual (ontogeny) and the history of the group to which it beiongs (phylogeny). Using Haeckel’s nomenclature, the three periods of ontogenesis, Anaplasis, Metaplasis, and Cataplasis, correlate with the three periods of phylogenesis, Epacme, Acme, and Paraeme. In addition to this general correlation, we now find that during the epacme of a group the development of individuals is anaplastic or progressively direct ; during the acme of a group, metaplastic or progressively indirect; and during the paracme of a group, cataplastic, or retrogressively direct. We have also found, that, in the history even of small groups, the epacme, acme, and paracme may often succeed one another in geologic time, and show similar correlations, so that we can often distinguish epacmic faunas, acmic faunas, and paracmic faunas in chronological succession. In series, also, epacmic forms, acmic forms, and paracmic forms, either in series of species or varieties, may occur in geological succession in different faunas, or in zoélogical grada- tion in the same fauna. 18. Law of Variation.— The action of physical changes takes effect upon an irritable, plastic organism, which necessarily responds to external stimulant by an internal reaction or effort. This action from within upon the parts of organisms modifies their heredi- tary forms by the production of new growths or changes, which are, therefore, adapted or suitable to the conditions of the habitat, and are therefore physiologically and organically equivalent to the physical agents and forces from which they directly or indirectly origi- nated. In so far, then, as causes and habits are similar, they probably produce representa- tion or morphological equivalence in different series of the same type in similar habitats ; and in so far as they are different, they probably produce the differentials which distin- euish series and groups from each other. PREFACE. DSi 19. The radical and epacmic forms of the Arietide probably originated in the North- eastern Alps, and migrated from thence southerly and westerly into Italy, and also in another direction westerly into South Germany and the Cédte-d’Or. In these last two faunas new series of acmic forms arose by modification, and these and the paracmic forms which seem to have arisen in the same basins flowed back into the Northeastern Alps, and thence into Italy, during Bucklandian and later times. They were also dis- tributed from these two basins to all others to the north and south of them in Central Europe. The Northeastern Alps and the South German and Céte-d’Or basins constitute a Zone of Autochthones for the Arietide, and other faunas to the north and south of these are what we have called Residual Faunas. The materials in the Museum of Comparative Zodlogy consist of various collections made in England by Damon, Marder, and Wright; Boucault’s famous collection from the Cote-d’Or, containing several of D’Orbigny’s types, and in part named by him, or by direct comparison with his collection; a special and very large general collection, especially rich, however, in South German species, purchased from Dr. Krantz; a valuable exchange from the Museum at Stuttgardt named by Professor Fraas; Professor Bronn’s collection labelled by him; a number of valuable species, principally from Belgium, from L. de Koninck’s collection; a similar lot presented by Prof. J. Marcou, from various localities in Europe; and others not sufficiently important to be men- tioned here. ALPHEUS HYATT. CamBripeGE, April, 1889. GENESIS OF THE ARIETID A. i INTRODUCTION. ORIGIN AND CHARACTERISTICS OF SUBORDERS. HE succession of forms among the silurian members of the genus Mimoceras indicates that true gyroceran shells occurred among Ammonoidea, differing from the similar forms among Nautiloids only in the possession of a globular pro- toconch and asmall ventral lobe. In some silurian and devonian Anarcestes these permanent adult stages are repeated in the development of the young. Those in Minoceras compressum are truly eyrtoceran, or open curves at first; and in others, as in a variety of Anarcestes fecundus described by Barrande, they are straight. The next stage of growth is a loose-coiled or gyroceran form, like the adult of Mimoceras. These stages can only be accounted for as hereditary tendencies of growth in a type which is being rapidly changed from a primitive ancestral straight form with simple sutures into a close-coiled nautilian shell. Branco” describes and figures a specimen of Bactrites with a protoconch similar to the very peculiar ovoid protoconch of Mim. compressum. He quotes Beyrich, who gave him this specimen, as authority for the view that Bactrites is connected with Mimoceras as Baculites is with the normal Ammonoids of the Cretaceous. This idea was first published by Quenstedt in his “ Die Cephalo- poden,”’ and it is quite possible that Bactrites of the Devonian may be a de- graded form of Mimoceras, but in that case the latter is also a degraded form of Anarcestes, or transitional between it and Bactrites. To establish this proposi- tion, forms of Mimoceras and Anarcestes should be produced in which uncoiling occurred in adults after a close-coiled stage of growth had been passed through. Such degraded forms are common in the Jura and Cretaceous. and enable the observer to connect Baculites with the normal coiled Ammonoids of the same formations. Whether this be so or not, the straight Bactrites-like young of some forms of Anarcestes, the gyroceran young of others of the Goniatitine, and the gyroceran adults and young of Mimoceras, indicate the derivation of Goniatitinee to have been from silurian straight shells similar to Bactrites, if not directly from that genus itself. 1 Genera Foss. Ceph., pp. 803, 304, 509, Proc. Bost. Soc. Nat. Hist., XXII., 1883. 2 Zeitsch. Deutsch. Geol. Gesell., XX XVII. p. 1. 1 2 GENESIS OF THE ARIETIDA. We pointed out in “ Embryology of Fossil Cephalopods,”* that the loosely coiled stages prevalent among Nautilinidae were repeated in the early stages of development in some of the Goniatitinge and in the later Ammonoids. This repetition was indicated by the form of the embryo which was flattened and depressed, and also in the first sutures and in the embryonal umbilici. These last are two conical or flattened depressions on either side of the protoconch, at its Junction with the apex of the conch. ‘They were accounted for as remnants of the umbilical perforation found in the young and adults of Mimoceras and all coiled Nautiloids. In our “Genera of Fossil Cephalopods” we narrowed this generalization by comparing the first whorl of the embryo in the close-coiled Goniatitinse and in all Ammonitinse with Anarcestes, thus bringing the affinities of all the Ammonoidea to a focus in the silurian genus Anarcestes. These and other similar observations, published before and since the work quoted above, have been founded upon the law of acceleration formulated also im the Preface of this monograph, pp. v, vi, Art. 11 and 14. Dr. Branco’s extensive and accurate researches? have shown that all of these opinions, though founded upon.a few specimens only, were sound, and that the law of acceleration can be relied upon as-a working hypothesis. Though treating us otherwise with more than just appreciation, this author failed to notice that we had used the law of acceleration in development, or made our inductions with the view of demonstrating its truth as a working hypothesis, and consequently attributed the discovery of this law to Wiirtenberger. Among Nautiloids the straight shells in each series appeared first ; they were succeeded by the cyrtoceran, gyroceran, and close-coiled. Among Ammonoids there is only one series — Bactrites, Mimoceras, and Anarcestes — which is parallel with any one series of the many occurring among Nautiloids. The open-whorled stages of the young of Anarcestes and other Goniatitine repre- sent a transitional and highly accelerated development. This transitional character is also indicated by the fact that, except in Mimoceras and some species of Anarcestes, the occurrence of the gyroceran form, even in the young, is sporadic. It occurs, as demonstrated by Barrande, in one variety of Gym- niles fecundus, and not in the other. Sandberger has shown similar though less marked variations in the young of Azar. subnautilinus, and Branco has described the embryo of var. viftiger of the same species as close-coiled. Other examples might be given, but it only remains to notice Branco’s doubts of the accuracy of our drawings of the young of Gon. atratus and Gon. Listeri. Both of these were found by him to belong to his close-coiled division of the Asellati of the Carboniferous. Our drawings were made with a camera. The details they contain show, better than any defence we can make, that they were also closely studied by the author, and often corrected before being placed upon stone. They indicate that primitive gyroceran forms of young are occasionally found even among the highest forms of carboniferous Goniatitine. 1 See especially articles ‘‘ Whorls” and “ Umbilicus,”” Bull. Mus. Comp. Zool., III. No. 5. 2 Paleontogr., 1880, 1881, XXVI., XX VII. ~— ORIGIN AND CHARACTERISTICS OF SUBORDERS. OS The depressed semi-lunar whorl appears first in the adults of Anarcestes. It is subsequently found in the young as a stage immediately succeeding the more cylindrical whorl of the gyroceran stage, when that occurs. In very close-coiled forms, the latter may be omitted, or be only slightly indicated, and then the anarcestian whorl appears at the beginning of the apex. In fact, this tendency in Latisellati, and especially in Angustisellati, affects the shape of the protoconch which is excessively depressed in the embryos of the higher suborders. We have, therefore, considered it convenient to designate the anarces- tian form of whorl as the primary radical of the Ammonoidea, reserving the terms primitive and transitional radicals for the straight and gyroceran modi- ‘fications as they appear in Bactrites and Mimoceras. The different series of the Clymeninz and Goniatitinee, and the Arcestins, often begin with, and maintain persistently in full-grown shells, the primary radical form. The Ceratitinze, Lytoceratinz, and Ammonitine, on the contrary, have this depressed form but rarely, except in their protoconchial stage, — and at the beginning of the apex or true conch, while it remains in what we have called the goniatitic stage of development. The Clymenine of the Devonian begin, when zoblogically arranged, with discoidal forms haying depressed semi-lunar anarcestian whorls. These de- pressed whorls are exchanged in the higher forms for compressed discoidal whorls, and these in turn for compressed involute whorls. The suborder includes several genera and in each there occur examples of this mode of succession, or rather procession, of forms, forming parallel series.! The sutures of the genera Beneckia, Longobardites, Lecanites, Norites, Meekoceras, Hungarites, and Carnites show them to be true Ceratitine, We should, with our present information, be disposed to include these, and all the genera mentioned by Mojsisovics as belonging to his group of Ammonites trachyostraca, in the Ceratitinse, distinguishing them by their well-known and peculiar sutures from the Arcestine, Ammonitine, and Goniatitine. The more or less compressed whorl, which in section can be described as helmet-shaped, is the natural successor of the depressed anarcestian whorl both in the growth of individuals and in the evolution of series of species. We have considered this in the work quoted, therefore, as the secondary radical. The secondary radicals” are prevalent in the Ceratitine, as shown by the extensive researches of Mojsisovics in the remarkable and masterly treatise above quoted. They completely replace the primary radicals as generators of series in the Trias, except in the paleozoic survivors of the suborder Arcestine. So far as the sutures are concerned, however, the Ceratitine, though distinctly characteristic of the triassic fauns, are like the Goniatitine. The young of Longobardites is really a Goniatite, similar to Prolecanites. 1 Genera of Fossil Cephalopods, Proc. Bost. Soc. Nat. Hist., XXII. p. 312. 2 We formerly included (Gen. Foss. Ceph., p. 324) in secondary radicals some quadragonal whorls like those of the adults of Xenodiscus; but we are now disposed to consider this an error, arising from not hay- ing observed that the young of these forms often possessed, during earlier stages of growth, the secondary or helmet-shaped whorl. This evidence shows that, in the most ancient periods as well as in later times, quadragonal whorls were derivative modifications of the compressed helmet-shaped secondary radicals. 4 GENESIS OF THE ARIETIDA. Norites is considered by Mojsisovies as allied to Pronorites, a genus of Gonia- titine, and by Griesbach, Zittel, and the author as allied more nearly to another genus of the same suborder, Sageceras. Throughout the group the lobes and saddles form a simple series im which very little differentiation is observable except in the highest forms. The ventral lobe is very broad and short, and the siphonal saddle broad and shallow. The survival of prolecanitian characters in these outlines is apparent the moment we dispense with the denticulations of the lobes and reduce the sutures to their primitive outlines. The Arcestins of the Dyas are known only by one species, described by Waagen, Cyclolobus Oldhami,' which has whorls of the anarcestian shape. It is an involute species, and there may be others of this genus in the same formation, not yet discovered, which have more discoidal whorls. According to our mode of translating the affinities of the forms, they arrange themselves as follows. Popanoceras of the Dyas, as the direct descendant of Prolecanites, inherits the tendency to have lobes and saddles of very nearly the same size, with lobes having trifid or bifid terminations similar to those of the young of Monophyllites, and also transitional to the sutures of the dyassic Cyclolobus, the most ancient of the true Arcestine. If we are right, the young of this last form, when examined, will be found to be similar to Popanoceras aniquum at a stage when its sutures have not yet acquired marginal lobes. The siphonal saddle in these forms and in true Arcestinge is small, often attenuated, and the ventral lobe large and often broad. The remaining lobes and saddles are more nearly of the same size, numerous, and formed a gradually lessening series mclining towards the umbilicus. The same aspect is common in the simpler shells of Megaphyllites and Monophyllites, but in these the large phylliform saddles, with entire outlines at their bases, exhibit closer approach to the Prolecanitidz. Arcestine, therefore, retain in their sutures the proportions of paleozoic forms of Goniatitine which have numerous lobes, but depart from them in having more complicated and ornate marginal digitations. The series, with some exceptions, have involved whorls which can only be considered as parallel with the more involute shells of silurian and devonian Anarcestes. With respect to its forms and the smoothness of the shell this series is a survival of purely paleozoic modifications. The Lytoceratine form a separate phylum, distinguished usually by the absence of true pile (ribs), the larval form and characteristics of the adult shell, and the leaf-shaped marginal saddles of the sutures. lLytoceras, in its smooth or unpilated shell, rounded abdomen, peculiar siphonal saddle, and phylliform marginal saddles, appears to be a more progressive form of the same genetic series as Megaphyllites and Monophyllites of the Trias. Even the peculiar coarse striations of the shells of these genera are often repeated among the Lytoceratinse of the Jura. Megaphyllites of the Trias is evidently closely allied to Monophyllites. The siphonal -saddle is similar to that of Monophyllites, and the marginal 1 Arcestes priscus, Waagen, is probably also a species of Cyclolobus. Geol. Sury. Ind., Salt Range, ser, 13, I. i, pl. ii. fig. 6. ORIGIN AND CHARACTERISTICS OF SUBORDERS. 5 saddles are phylliform, The young of Monophyllites Suess?, Moj.,' of the Trias, has sutures similar to the adults of Popanoceras antiquum? and Kingianum of the Dyas,? which are true Goniatitine. The sutures of Popanoceras are in their turn transitional between Monophyllites and the more normal Goniatitine of the genus Prolecanites. Triassic Ammonoidea have shallow ventral lobes and very prominent broad siphonal saddles, thus giving the first lateral saddles the aspect of being ad- juncts of the siphonal saddle. In consequence of the more direct descent of Lytoceratine of the Jura from primitive forms, their sutures persist in retaining triassic outlines, having usually short abdominal lobes, large siphonal saddles, with the superior laterals apparently set upon their sides, the larger lobes expanded and profusely branching at the top, the saddles expanded and profusely branching at the base, the auxiliary lobes and saddles more numerous and more nearly equal to the larger lobes and saddles than in Ammonitine. Neumayer has demonstrated trumpet-like apertures in Lyf. im- mane.* 'The frilled and elevated ridges in shells of many forms indicate that these are perhaps not uncommon in this group.’ The normal forms of the Ammonitinze, the Arietidee of the Lower Lias, ean be united to the genus Gymnites through Psiloceras. Gymnites can be traced back to the Goniatitinz through Arcestes of the Trias and Cyclolobus of the Dyas. The Ammonitinz do not, therefore, come directly from the Goniatitinze, as do the Lytoceratinsz, but are probably direct offshoots of the lower Arcestine. The Ammonitine include not only the typical jurassic and cretaceous forms, but also the allied radical genera Schlotheimia and Psilo- ceras of the Lias, and Gymnites and Ptychites of the Trias.° In Gymnites of the Trias, the primary radical is exchanged for the more compressed discoidal secondary radical, but still smooth shell, which is also characteristic of Psiloceras of the Lias. The sutures are correlatively modified, and begin to assume the aspect and proportions of the true Ammonitine. The siphonal saddle is more prominent, but still retains in many species the pointed aspect derived from the Goniatitinze. The narrow first lateral saddles are apt to appear like adjuncts of the siphonal saddle, owing to the great size and breadth 1 Mediterr. Triasprov., pl. Ixxix. fig. + a-e. 2 Arcestes antiquus, Waagen, Salt Range, Pal. Ind., ser. 13, I. i, pl. i. fig. 10. 8 Russia and Ural, M. V. K., II. pl. xxvii. fic. 5. 4 Mojsis. et Neum., Beitr., IIT., 1583, 1884, pl. xx. 5 Schlonbach, Paleontogr., XIII. p. 169, pl. xxvii. fig. 3, deseribes Amm. hircicornis, one of the Lytoceratine, having a series of prominent flaring ridges indicating permanent apertures of similar form. The slight, blunt rostrum is a notable characteristic of these apertures. Unfortunately, very few have been preserved, possibly owing to the fact that they were in most species, as in the two mentioned, thin flaring ridges, easily destroyed. We can only suggest, therefore, that this form of rostrum might have been peculiar to this suborder, 6 In “ Genera of Fossil Cephalopods,” in 1883, we expressed this opinion as follows: ‘‘This genus (Cyclolobus) is very important, since it enables us to show the gradations by which the Prolecanitide approximate to Arcestes, Ptychites, and Monophyllites.’? Mojsisovies, with new materials from Spitz- bergen, has lately demonstrated the correctness of this opinion in part, and gives conclusive evidence of the probable derivation of Arcestes and Ptychites from Popanoceras. Arkt. Trias Fauna, Mem. Akad. St. Petersb., XX XIII. No. 6, p. 66, pl. xv. 6 GENESIS OF THE ARIETIDA. of the first lateral lobes and the shortness of the ventral lobe. The aspect of the second laterals in many species, and the gradation from these into the auxiliary lobes, show that they retain the more primitive aspect of the earlier forms in this part of the sutures. . Ptychites of the Trias has sutures similar to those of Gymnites, and the modified aspect of marginal lobes and saddles in both genera shows that, in spite of a near approach or resemblance in the sutures to many Lytoceratine, they cannot be considered as so nearly related to them as to Cyclolobus. Mojsisovics says that the evidence of genetic connection of Pséloceras planorbe and Gymuites incultus vests alone upon the resemblances of the auxiliary lobes and saddles, and that the resemblances in form only occur between the discoidal Gymnites and the most involute Psiloceratites, the former being indeed much more involute than the most involute of the Psiloceratites. The genus Halorites of the Trias is regarded by Mojsisovies as the probable ancestor of the Arietidee. We cannot recognize that there are any very marked differences in the amount of involution or form between Gym. incultus and Gym. Paina when compared with Psi. planorbe, and the resemblances of the sutures are exceed- ingly close, especially when the species of Psiloceras of the Mediterranean province are studied. The aspect of the shells im the three former are very similar, while in the types of Halorites already cited by Mojsisovies, Hal. Ramsaueri, semiplicatus, decrescens, and semiglobosus,, they are very distinct. The range of form in Halorites embraces highly sculptured shells, altogether triassic in aspect. | Neumayr’s? and Wiihner’s® researches entirely confirm the position here taken and show that Psiloceras possessed a series of involute shells. Psiloceras and Gymnites, therefore, appear to be two parallel genera of the same group, in each of which discoidal forms give rise to more involute shells. Gym. incultus may be traced into the more involute Gym. Humboldti, and the still more involute Gym. Credneri. The adolescent young of Gym. Paina, Mojsis.* and tncultus® show less involution than the adult, and we may confidently expect that some correspondingly still less involute discoidal ancestral forms will be found. Mojsisovics has not yet published his observations in full, and his evidence is therefore not completed; but, so far as we now know, the deriva- tion of Psiloceras seems to have been from Gymnites as a common ancestor and not from any forms of the Ceratitinze like Halorites or its allies. Mojsisovics has said, that out of his group of Ammonites leiostraca the genus Phylloceras alone persists and is but little changed in the Jura; whereas the Anm. trachyostraca, or Ceratitinse, are more largely perpetuated, though much changed, in the true Ammonitine. Our view differs, since we consider all groups of the Trias to have been discontinued in the Jura except the Lytocera- tine. Itis probable that a close affinity existed between Psiloceras and Gym- nites, and the former is a modified Triassic survivor in the Lias; but the constant reappearance of the psiloceran form in the young of undoubted Arietian Amm. Gattungen, Verhand. Geol. Reichs., 1879, No. 7. 1 2 Unterst. Lias, Abhandl., Geol. Reich., VII. 8 Unt. Lias, Mojsis. et Neum , Beitr., III. 4 Med. Triasproyv., pl. lvii. fig. 2. 5 Thid., pl. liv. fig. 3. ORIGIN AND CHARACTERISTICS OF SUBORDERS. 7 species shows that we must reckon it among the Arietide. The genus Schlo- theimia is also a purely jurassic series, though undoubtedly triassic in respect to its sutures. The young of Schlotheimia calenata is an almost. exact reproduction of the form described by Mojsisovics as Ayoceras Buonarotti in “ Jahrbuch Geologi- schen Reichsanstalt,’ 1 and afterwards referred to Celtites in his “ Mediterranean Triasprovinz.”* The pile cross the whorls on the abdomen in the same way, and the general aspect of this discoidal shell is similar. It seems quite likely that this is a young shell of some species, and until its exact affinities can be determined it is of no great value. At present it would be difficult to say with any certainty to what genus it might be referred. Mojsisovies was evi- dently in doubt, since he states that it may be a young form of some species of Balatonites. The resemblance to the young of Schilot. catenata may be due to a purely pathological deformation, since the crossing of the abdomen by the pile occurs from disease in many species of the Arietidse: and other keeled groups of the Jura, notwithstanding the fact that it is normal in others. These facts, and the gradations of form between Schlotheimia and Psiloceras presented by the genus Waehneroceras,’ and by the young of this last genus, lead us to think that Schlotheimia was derived from Psiloceras. The Ammonitine of the Jura, so far as known, show no special traces of theif prolecantian descent, except in the discoidal shells and phylliform sutures of the genera just mentioned, and in the embryonic and generaiized goniatitic characters of the apical stages of the shell. The ventral lobe of the Ammonitine is deep and narrow, the siphonal saddle small but more or less dentated by marginal lobes and saddles. The lateral saddles are broad and not so deeply divided by marginal lobes as in the Lytoceratins, the lobes are narrower at the tops than in that suborder, and the saddles consequently narrower at their bases. The great size and small number of the lobes is also a marked peculiarity. The superior lateral saddles and lobes are especially remarkable for size, and the auxiliary lobes and saddles much less important and more unequal as compared with the lateral lobes and saddles than in Lytoceratinze. The marginal lobes and saddles are as a rule short and pointed, and the saddles rounded, but not phylliform. Possibly another distinction will eventually be demonstrated in the more constricted and rostrated apertures of many of the Ammonitine. The characteristics of the embryos and of the earliest stages do not yet seem sufficiently well known to be used in this connection. The Ammonoids, therefore, according to our views, are not divisible into two grand divisions, but have six suborders: the Goniatitine, of the Silurian, Devonian, Carboniferous, Dyas, and Trias; the Clymenine of the Devonian ; the Arcestinzs of the Dyas and Trias; the Ceratitine of the Dyas and Trias ; the Lytoceratinee of the Trias, Jura, and Cretaceous; and the Ammonitine of the Trias, Jura, and Cretaceous. 2 Unfortunately, there is not space enough within the necessary limits of this monograph to discuss the classifications of Mojsisovies, Fischer, and Zittel, 1 Vol. XIX., 1869, pl. xv. 2 Page 129, pl. xxix. ® A new genus described in this memoir. 8 GENESIS OF THE ARIETIDA. and the embryological divisions proposed “by Branco.' The classification given above was necessary in order to introduce our remarks upon the Am- monitine, and show clearly why wé limited this suborder as defined above ; any further discussion would lead us too far away from the immediate objects of this memoir. NOMENCLATURE OF STAGES oF GrowTH AND DECLINE. In a paper read before the Boston Society of Natural History, November 16, 1887, the author discussed the classification of the stages of growth and decline, dividing them as follows : — 1. The earlier stages, embracing the ovum (monoplast, Lankester), the monoplacula, and the diploplacula, were considered under one term, Protembryo, because of their parallelisms with the single and colonial Protozoa. 2. The next, or blastula stages, were classified under the head of Mesem- bryo, on account of their resemblances to the Mesozoa; the latter being those forms usually included in the sub-kingdom of Protozoa, but which have true ova and spermatozoa, and can be therefore separated as one-layered, spherical Blastrea, closely parallel with tlie blastula, and precisely intermediate between Protozoa and Metazoa. 3. The gastrula stages were considered as referable to true Metazoa, and were styled accordingly the Metembryo. 4. The earlier planula or ciliated stages were regarded as indicating a still very remote ancestral type, in common with Semper, Lankester, and Balfour, and were termed the Neoembryo. 5. The later ciliated stages— those which show the essential characters of the type to which the embryos belong — were classified as the Typembryo; ex. the veliger, nauplius, etc. The typembryos were considered as the last of embryonic stages, and those which followed were regarded as true larve on account of their more demonstrable connections with well known forms. It was found by applying this classification to the fossil Cephalopoda that the pro- toconch of Owen was the shell of a univalve typembryo, which must have been a veliger not very widely removed in structure from the similar shells of the embryos of Gasteropoda and Pteropoda.* The principal difficulty of the application of this view lies in bringing the wrinkled and curious forms which occur upon the apices of some Nautiloids into 1 Mojsisovics, Med. Triasprovinz; Fischer, Manuel de Conchyliologie; Zittel, Handbuch der Paleon- tologie; Branco, Paleontogr., XXVI., XXVII. 2 Robert Tracy Jackson, a pupil of the author, in an essay now in preparation (‘‘ Phylogeny of the Pelyeypoda’’), shows that the typembryo stage of mollusks is limited to an early period characterized by the existence of a shell-gland and the plate-like beginnings of a shell. Later veliger stages, he says, are ref- erable to the class or phylum of Mollusca. to which the embryo really belongs, and he names them ‘‘ Phyl- embryo”’ stages. The ‘‘ prodissoconch”’ is a name given by Jackson to the embryonic, bivalvular shell of Pelyeypoda, which is the equivalent of the protoconch of cephalous mollusca. The completed protoconch of the cephalous mollusea, and prodissoconch of Pelyeypoda, Jackson considers as a stage later than that at which the phylembryonic characters are emphasized, and as the close of the embryonic shell period. His paper will give types of these and other stages considered in the several classes of mollusks. NOMENCLATURE OF STAGES OF GROWTH AND DECLINE. 9 exact relations with the indubitable protoconchs occurring upon the apices of the conchs of Ammonoids and Belemnoids. The wrinkled lump above referred to is unquestionably a part of the shell. It is not only closely attached, but the longi- tudinal striz of the apex of the true conch are continuous upon the proximate parts of the lump. It had an aperture which must have remained open until the body of the veliger had entirely left the interior of the protoconch, and was then closed by the apical plate. There is a cicatrix upon the apex of the conch, which is invariably concealed by the lump when it is present, and in some examples we observed the fracture of the outer layer of the shell on the apex of the conch and outside of the ordinary boundary of the cicatrix, which could only have been caused by the violent removal of the lump. The wrinkled and contracted aspect of the lump when preserved can be accounted for by assuming it to have been composed of conchiolin. This also accounts for its almost invari- able absence, since such an organ must have been easily lost or destroyed. The lumps must consequently be regarded as the remnants of conchiolinous pro- toconchs having elongated and narrow apertures; but probably they were, when in a living condition, much larger and more oval, and more similar to the protoconch of the Ammonoids. The continuity of the striae from the conch to the protoconch also shows that the conch was built out from the aperture of the protoconch, layer after layer, and the concentric markings, and form of the apex, which correlates with that of the scar, sustain this idea. The figures on the fol- lowing pages are less perfect than several other examples studied by the author since these were drawn. They do not show the passage of the external longitu- dinal striz from the apex of the conch on to the surface of the protoconch. A living chamber among recent and fossil Nautiloids marked a period of rest after a stage of growth. The septum, therefore, was not built until the animal arrived near the final steps, or had altogether stopped building out the sides of that part of the shell in which it lived. At any rate, we can say with- out risk of error that the septum was the final step, or one of the final steps, in the construction of a living chamber. 6. The first living chamber, or the first larval or nzpionic! stage of a Nauti- loid was, therefore, represented by the apex of the conch in that order; but the first septum and siphonal czecum did not exist at this stage, which is represented by a straight or slightly curved widely spreading cone, — in fact, the empty apex of the conch. The length of the first living chamber has not been ascertained ; but that it was short seems probable from the form of the cone in Nautilus. Doubt- less this remark does not apply to the earliest forms of closely coiled shells, in which the cone was much slenderer than in existing Nautilus, and the first living 1 Nymws, infant, or young animal. The term ‘ silphologic ’’ was used for this stace in the article above quoted. ‘This literally means ‘ grub”? stage, and it is not strictly applicable to a normal progressive stage of development. Grubs, caterpillars, and the like, among insects, are degraded or retrogressive develop- ments, as compared with the normal, probably hereditary Thysanuriform lary of what are commonly called the lower orders of Insecta. Studies of insects lately made have convinced us of the truth of this opinion, first published by Friedrich Brauer, and of the need of changing this term to the one used, and of reserving silphologic as a general term for retrogressive stages, such as one finds in the larye of Coleoptera, Lepidotera, Hymenoptera, and Neuroptera, 9 10 GENESIS OF THE ARIETID As. chamber perhaps longer in proportion. The entire absence of a cecum, and of all signs of a siphon, may be inferred with probable certainty in this first stage ; and we proposed, in the paper referred to above, to name it the Asiphonula. This form may indicate the previous existence of a common univalve ancestor for the Cephalopoda which resembled the Pteropoda. Certainly the aspect of the calcare- ous protoconch of Ammonoids and Belemmoids favors this idea, first suggested by Von Jhering ; and the asiphonula adds another argument, since it has no siphon or true septum. The young of the Pteropoda, especially the ancient forms, had calcareous protoconchs in most forms; but doubtless there are more primitive shells in which the protoconchs had the more primitive, embryonic, conchioli- nous stage of development.. The evidence, therefore, is not conclu- sive, but it justifies the supposition that Cephalopods and Pteropods had originally some common an- cestor, a true shell without septa or siphon, and possessing a proto- conch, which might have been conchiolinous. There is, however, another group, the Scaphopoda, which may claim to be considered in this connection. According to W. K. Brooks, the veliger is rep- resented by the adult of Dentalium Fig. 1-3. Apex and protoconch of Orth. elegans, Miinst., Seam in several of its leading charac- te sie, below and infront. In Fig 2 the he aie relly tovisties, and this must be regarded conch (a). Named by Klipstein, Loc. St. Cassian, Coll. Brit- therefore as the most generalized ish Museum. Fig. 4,5. Apex and protoconch of another specimen mounted type of the true Mollusca. It is with the first on the same card. Named by same, Loc. same, Coll. same. quite possible that the asiphonula Fig. 6-8. Views from the side, front, and below of the same parts in Orth. politum, Klipst. ‘The shading on the protoconch may have retained some of the of Fig. 8 does not indicate structure; this protuberance is : smooth. a, protoconch; b, shoulder of the area of the cica- characters of the veliger, and may trix. Named by same, Loc. same, Coll. same. here resembled Dentalinnnaen some common ancestor, and may have descended from this form without having passed through any pteropod-like ancestral modification. The peculiar resem- blances of the young of some of the Goniatitinee and the adults of Tentaculites among Pteropoda may be entirely due to homoplasy, and not to homogeny.* 1 These terms were first used by Lankester (Jour. Micr. Sci., XVII., 1877, p. 486). They express phenomena with which naturalists have long been familiar, ‘‘homoplastic ’? meaning representation and independent origin of similar characters, and ‘* homogenous ’’ meaning genetic connection. See also previ- ous use of terms Heterology and Homology for the same phenomena by Cope, in his masterly essay, ‘* Origin of Genera,” Proc. Acad. Sci. Phila., 1868, and “ Origin of the Fittest,’”’ p. 95. ee aa = NOMENCLATURE OF STAGES OF GROWTH AND DECLINE. 11 It is obvious that we cannot ac- count for the nautilus-like ven- tral saddle of the earlier sutures of the Ammonoids, the calca- reous shell of the protoconch, the coecal stave, the absence of the collar in the lower Goniati- tine and in the young of the higher forms, the often central position of the siphon in the young, and many other charac- ters, unless we admit a proba- ble derivation of the Goniatitinz from some straight microsipho- nulate form of Nautiloid. It is, therefore, highly probable that the pteropod-like aspect of the youne of some Goniatitinee may be a purely homoplastic charac- ter, and be meaningless so far as the genesis of the group is con- cerned. 7. The next or second of the nepionic stages was represented by a living chamber, which was completed by the building of the first septum with its attached excum, indicating the primitive beginnings of a siphon. This stage we styled the Czcosipho- nula, and we have considered the possession of a cxcum to be an indication of the former exist- ence of an ancestor having a central series of caecal pouches. These may haye had functional communication in some forms by means of an endosiphon, as in the Endoceratide, and in others, either belonging to this family or to a more primitive group, they may have been closed cxca. 8. The next nepionic stage was ended when the second sep- tum was built in the modern Fig. 9,10. Views from the side and below of the plug which the animal of Orth. truncatum, Barr., habitually built on the exterior of the broken or truncated end of its shell. The last suture is shown in Fig. 9, and the internal shadowy markings are apparent in both figures at a,g. These, however, in Fig. 9, are too far removed from the exterior. When the outer layer of the plug is penetrated, they are seen to be a part of its structure. ‘The side view is also defective in the drawing of the pseudo siphon (d). There should be three distinct steps indicating three layers. ‘The external crenulated striae of the plug appear at h. Loc. Bohemia, Coll. British Museum. Fig. 11, 12. Views of the same from the side and below, to show the external markings of the plug (h), which contrast strongly with the perfectly smooth shell above the septum of trunca- tion and internal strie (1) which appear when the outer layer is fractured No septa ever occur in the plugs. These figures are introduced in order to meet M. Barrande’s objections (Syst. Syl, Pl. 488), that the examples of what we have called the protoconch and apex of the true conch were in reality plugs similar to those of Orth. truncatum. There is no need of mak- ing any remarks; if our figures are correct, we are right in our statements. It may, however, be well in this connection to say that M. Barrande has done us the honor to make use of a number of our figures, including in part the above. Fig. 18-15. Views of the cicatrix and apex of Orth. unguis, Phill., after the shedding or remoyal of the protoconch as it usually occurs, leaving the cicatrix uninjured. Fig. 13 shows the area of the cicatrix much enlarged; b, conch or apex forming a smooth shoulder; and ec, depressed surface of- the cicatrix. Fig. 14, view less magnified of apex; Fig. 15, section of same. Toe. Dublin, Coll. British Museum. Fig. 16. Apex of Orth. unguis, Phill., natural size, with first three sutures. Fig. 17. Apex of same species after the probably violent removal of the protoconch, showing the fractured shell (b), and the unusual aspect of the cicatrix. ‘This and Fig. 16 are types. Loc. Yorkshire, Coll. British Museum. Fig. 18. Front view of Fig. 19. The broken line (k) is hypotheti- cal. It indicates the possible outline and position of the cecum, supposing the oval area in the centre of Fig. 19 to have repre- sented that organ. Fig. 19. Apex of Orth. politum from below. The protoconch has also been yiolently removed, and the opening plugged, apparently from within. ‘The dark spot on the right seemed to be a rup- ture in the external surface. ‘The oval shade in the centre indi- cated an internal structure, which may have been the caecum in the first air-chamber. Loe. St. Cassian, Coll. British Museum, 0G 12 GENESIS OF THE ARIETIDA. Nautilus, and in the vast majority of all known fossils of the order so far known to the author this stage had similar characters. The siphon was larger at this stage than subsequently, and possessed a prolongation which reached down into and lined the primitive cecum. This closed pipe was however more or less cylin- drical, and formed a transition to a cylindrical, open, siphonal tube, when com- pared with the cecum on the one hand and the siphon of the succeeding septa on the other. The second septum was prolonged apically into a funnel, and this was continuous with a true porous wall, which formed the remainder of the pouch. We have already pointed out the probability that this wall was the homologue of the calcareous sheaths or endocones which filled the imteriors of the siphons of Endoceratide. There is, therefore, as previously stated by the author, a structural though highly concentrated and much modified remnant of the adult siphonal elements of an Endoceras still preserved in this stage, even in the existing Nautilus, and we propose to name it the Macrosiphonula. 9. The next nepionic stage in living Nautilus was the third living chamber and third septum with its siphon. The siphon has a true funnel, and the siphonal wall attached to it is less swollen out, and seems, upon re-examination of the junctions at the opening of the funnel in the second septum, to be discontinuous. If we are correct, this stage has a small siphon consisting of the usual funnel and tubular porous wall, as in the vast majority of all Nautiloidsand Ammonoids. We proposed to name this, according to its aspect and structure, the Microsiphonula. The microsiphonula, though a neepionic stage in the modern Nautilus, did not always occur among larval stages, but had in common with the macrosiphonula a traceable beginning in the adult stages of ancestral types. The genesis of the two forms of siphon may be studied in the Endoceratidee. In this family Cyrto- cerina had a siphon, which continually increased in size, probably throughout life, though more forms need to be described before one can be assured of this as a fact. There is no doubt, however, that the next form, Piloceras, had what we can safely call a macrosiphon of typical structure until very late in life. The large shells collected in Newfoundland by the author had siphons of great size, which were only slightly contracted or remotely approximated to the tubular condition of the Orthoceratide in the adolescent and adult stages. This and other changes occurring in the adolescent stages induced us to distinguish them by a special term, Nealogic. The adolescent or .nealogic stages, therefore, and the stages of the adult, or, as we have named them, the Ephebolic stages, in Piloceras show for the first time a tendency to contract the siphon or approximate to the microsiphonula, but they never had a true microsiphon. The contracted siphon in these forms, as in the other genera of this family, always had the holochoanoidal or complete funnel reaching from one septum to the next, and a series of conical concentric endocones, or sheaths, as they have been called by others, which stretched from the ends of the funnels, and were the homologues of the porous walls of the segments of the siphon in Nautilus. The terminations of the endocones were prolonged into a central tube, or endosiphon, which we have previously described, and which probably served as a NOMENCLATURE OF STAGES OF GROWTH AND DECLINE. 13 functional siphon in these shells. Gerard Holm! first called attention to the interesting character of the young stages of the siphon in Endoceras, and has shown this organ to have been very large even in the young, having not only a cecal beginning, as in other forms, but in several species having a swollen or macrosiphonulate form which endured throughout several septa. In speci- mens now in the Museum of Comparative Zodlogy, at least six septa were built before any signs of contraction began to appear. In other cases figured by Holm the siphonal cecum, though very large as compared with that of Orthoceras, was attached to the first septum, as in all the shells so far known from that group, and occupied only the first air-chamber. We should suggest to those having materials for study, that the shells having this last character are very likely not true Endoceratites, but perhaps the young of species of the genus Sannionites, which, according to the classification followed by the author, is a genus distinct from Endoceras, because the species possess a much slenderer siphon. Whatever the fate of this suggestion, it is plain that transitional series exist in this group between Sannionites and Cyrtocerina or Piloceras, and that gradations occurred also in Piloceras, which show that contraction of the siphon began first in adults, and then, according to the law of acceleration, was inherited in the nealogic stages of immediate descendants, and finally became nepionic in the smaller siphoned species of the genera Endoceras and Sannionites. This tendency to contraction in the diameter of the siphon indicated the beginning of a series of transformations which accompanied a decrease in size of the fleshy siphon, and other correlative transformations, such as the decrease in length of the funnels, and the contraction and straightening out of the calcareous endocones, so as to form the walls of a tubular siphon. In other words, as the siphon contracted, the func- tional endosiphon formed by the open and extended tips of the endocones was finally brought into line with the funnels, and together with them formed the microsiphon, which is consequently a degraded modification derived from the funnels, endosiphon, and endocones of the Endoceratidee. The Orthoceratidee and all the remaining forms, with some notable exceptions which we shall take up and describe in future papers, had a microsiphon. The whole microsiphon formed a continuous open tube of narrow diameter, reaching from the last septum to the nepionic septa, which represented the macrosiphonula. Doubtless the duration of nxpionic stages will be found to vary somewhat in ancient forms, but the indications, so far as known, are in favor of the theory that the vast majority of even ancient forms had a microsiphon, which was developed comparatively early in the life of the animal. The nealogic stages of succeeding groups would be very interesting if there were space to describe them, but we shall have to illustrate this part of our work among the Ammonitinz. The protoconchs of Ammonoidea, including the genus Bactrites, had, as remarked above, globose forms with calcareous shells, and these shells were continuous with the apex of the conch, but the aspect of the junctions was quite distinct from those of Nautiloids. The con- striction between them and the apex was very slight in the uncoiled young of the ? Dames et Kayser, Paleontol. Abhandl., IIT., Part I. 14 GENESIS OF THE ARIETIDZ. more primitive forms of several silurian and devonian species of Goniatitinge, and this is notably the case in Bactrites which has a straight shell. In these primitive forms the apertures of the protoconchs must have been less contracted than in most Nautiloids. The apex of the conch did not expand so fast as in Nautiloids, but was more nearly of the same diameter as the neck of the protoconch, and often remained tubular for a considerable portion of the nepionic period. ‘This was especially evident in the more open whorls of the anarcestian larvee, figured by Sandberger, Barrande, and Branco. Among the close-coiled forms of paleozoic species, and in still later occurring genera, the protoconch itself became depressed, and a deep dorsal constriction resulted from the abruptness with which the apical part of the conch turned in upon the inner (dorsal) side of the protoconch. The caleareous nature of the shell, the depressed form and transversely con- stricted aperture, and the closer union of protoconch and conch among Ammo- noids, separated the young apparently so widely from those of Nautiloids, as to lead Barrande, Munier-Chalmas, and Branco to deny that transitions occurred between them. Another distinction of importance was, that the ‘aperture of the protoconch was closed, not by an apical plate, but by the first septum. In other words, the asiphonula of Nautiloids disappeared as a distinct nzepionic stage, and the cexcosiphonula took its place in the development of the young among Ammonoids. This fact led Branco in his masterly work on the early stages to assert, in common with Barrande and Munier-Chalmas, that the protoconch of the Ammonoids was the homologue of the apex of the conch and first air-chamber in the Nautiloids. Certainly the calcareous shell and the position of the first sep- tum and czecum appear to be in favor of their view. On the other hand, the student of embryology will be slow to admit that the resemblances of the protoconch in Ammonoids to the veliger shell has no mean- ing. Ifit have any meaning at all, and can be compared with the protoconchs of the Cephalophora during the veliger stage, then during the whole of that stage the typembryos of Ammonoids, like all other veligers, could not have had a siphonal cecum or siphon. This is insured by the emptiness of the protoconch, the siphonal cecum being present only in the aperture, and not penetrating far back into or resting upon the first formed plate of the protoconch, as in the first air-chamber of Nautiloids, Another argument in favor of the view here advocated is the general fact, cited in the paper quoted above, upon the “ Values in Classification of the Stages of Growth and Decline,” that the typembryos, to which class of forms the veli- ger belongs, cannot be said to have the essential characters of any specialized division, like the Cephalopoda, but have to be compared with remote and gen- eralized types from whom their principal characteristics were inherited. The authors quoted above, holding the view that the protoconch was the homologue of the first chamber and apex of Nautiloids, necessarily rejected our theoretical explanation of the presence of the first septum and cecum in the aperture as due to acceleration of development. Nevertheless, this explanation still seems to us correct, and we have now a new a ii 6 i hs NOMENCLATURE OF STAGES OF GROWTH AND DECLINE. 15 r point to make in its favor. If the protoconch of Nautiloids was an empty con- chiolin shell and represented the veliger stage, it most certainly could not have been the ancestral form from which the calcareous tendency of the same stage in Ammonoids was derived. The characteristics of the asiphonula of Nautiloids are, however, just what are needed to fill the gap. The apex at this stage in Nau- tiloids is rounded and caleareous. The tendency to deposit calcareous matter could therefore have been inherited from an ancestor corresponding to the asi- phonula, and which we will name the Asiphonophora. The Asiphonophora must have had a calcareous shell acquired as an adaptive character, without internal calcareous septa or a siphon. ‘This form could not have been by any means so far removed from the ancestor of the veliger as the immediately following an- cestor of the macrosiphonula, which we have named the Macrosiphonophora. This must have had septa and a central axis of ceca, or at any rate at least one septum and a cecuni. The characters of the Asiphonophora, when transmitted to the Ammonoids according to the law of acceleration, would have been inherited earlier than in Nautiloids, would therefore have affected the growth of the protoconch, and would have necessarily produced the calcified shell of this stage in Ammonoids. The fusion of the protoconch with the conch in all Ammonoids was the imme- diate result of this process, and in this. way the more tubular form and freer connection of the protoconch with the true conch, and the constant adhesion of the former to the latter, can be explained. The disappearance of the asiphonula as a distinct stage in the young of the Ammonoids appears to us, therefore, not an argument against the deriva- tion of the Ammonoids from Asiphonophora, but in favor of this opinion. In fact, it seems to us that, in order to disprove it, opponents will have to find a cicatrix upon the apex of the protoconch in the Ammonoidea. According to the uncompromising attitude of those who insist upon the naked facts, and are hostile to explanations, the protoconch is the apex of the conch in Ammonoids, and the absence of any cicatrix upon the tip of this is a difficulty they can only surmount by asserting that the general and special homologies we have traced, and all the embryological and nzepiological correlations, are purely homoplastic, and do not indicate the derivation of the Ammonoids from any form of Nauti- loid. They must also explain away the similarity of the protoconch in external aspect to the veliger shell in Gasteropoda, since this is an earlier stage than that of the apex of the true conch in Nautiloids, Ammonoids, and all cephalous mollusks. Can any of these gentlemen tell us why the cicatrix does not appear upon the protoconch of Ammonoids, and explain at the same time how that shell came to be similar to the veliger shell in the Cephalophorous Mollusca on the one hand, and the apex of the conch of Nautilus on the other? It must be observed, also, that we do not insist that the primary radical of the Ammonoids, Anarcestes, was necessarily descended directly from Endoceras, but that it had probably come from a prototype like the veliger, possibly, as suggested by Brooks, from a class now only represented by the genus Dentalium. The next step, according to our translation of the evidences, must have been 16 GENESIS OF THE ARIETID. the Asiphonophora, which may have been more of a Pteropod or Scaphopod than a Cephalopod. So far as the shell goes, there are no similarities to the peculiar shell of Dentalium, but perhaps more to that of a Pteropod. The next step in this line of genesis must have been the ancestral generator of the characters of the czecosiphonula, which we propose to call the Cecophora, a form which must have been a reality in some shape, and in some species doubtless had the characters of the czecosiphonula in its ephebolic stages. This class of forms, though having septa and a central axis, which we might have to consider as a primitive siphon, was nevertheless quite distinct from those which followed. The next link in the genealogical tree must have been the ancestor of the peculiarities of the macrosiphonula, and this is luckily a well known form. The Endoceratidee enable us not only to see that the previous train of induc- tion is legitimate, but to connect our line of hypothetical forms with the next in the evolution of the group. The Endoceratide are true Macrosiphonophora, according to the nomenclature adopted here, and are transitional to the more highly specialized and stable modification which had what we have termed the microsiphon. When this organ came into being in the direct line of change, the evolution of the forms also changed its character, The more rapid or accelerated modes of change were replaced by slower processes. The changes occurring in the types preceding, and including the Hndoceratidz: (Macrosiphonophora), were, if we can judge by the abrupt transitions of the genera in this family, more rapid and more important in their effects on structures than was the rule subsequently. ‘This is also shown in the structural changes taking place in the embryos of Nautiloids and Ammonoids, as compared with the slow and comparatively slight changes of the subsequent stages of growth, The rapid acceleration of the macrosiphonulate character during the evolution of the Endoceratide, the still more rapid acceleration which took place in the evo- lution of the microsiphon among Ammonoids, and the fusion, through accelera- tion in development, of the characters of the asiphonula with the protoconch, all bear witness to the truth of this induction. The neepionic stages in ancient asellate forms of the Ammonitine, as has been shown above, may be considered as indicating the primitive radical, the straight orthoceran, and the gyroceran, or loosely coiled nautilian shells; but in 1 We have already traced the more rapid evolution of the ancient forms of Cephalopoda, and need not go into the matter any further in this monograph than to state that these facts accord with the law an- nounced in Genera of Fossil Cephalopods (Proc. Bost. Soc. Nat. Hist., XXII. p. 262), which reads as follows: ‘‘ These facts, and the acknowledged sudden appearance of all the distinct types of Invertebrata in the Paleozoic, and of the greater number of all existing and fossil types before the expiration of paleo- zoic time, speak strongly for the quicker evolution of forms in the Paleozoic, and indicate a general law of evolution. This, we think, can be formulated as follows: types are evolved more quickly, and exhibit greater structural differences between genetic groups of the same stock, while still near their point of origin than they do subsequently. The variations or differences may take place quickly in the fundamental structural charac- teristics, and even embryos may become different when in the earliest period, but subsequently only more superficial structures become subject to great variations.” See also Foss. Ceph. Mus. Comp., in Proc. Am. Assoc. Adv. Sci., XXXII, 1883, p. 338. NOMENCLATURE OF STAGES OF GROWTH AND DECLINE. ii all other forms, especially in the devonian latisellate and triassic angustisellate embryos, the tendency to become closely coiled, and to inherit the depressed primary radical whorl of Anarcestes, produced the Goniatitinula, and affected even the protoconch. The protoconch through heredity becomes depressed fusi- form by lateral expansion in the Angustisellati, and the embryonic nautiloid character of the first septum in the asellate forms and its tendency to form a broad ventral saddle in the latisellate and a narrow ventral saddle in the angustisellate embryo is correlative with this progression of form. The goniatitinula is a true larva, corresponding to adults within the order. We use the term because it is the characteristic larval form of the Ammonoidea, which was introduced at first among adult Goniatitine, and in the higher forms of this group became, by acceleration, fused with the microsiphonula. The remarkable researches of Branco enable us to state that this progres- sion in complication of the embryo in form and sutures has no counterpart in the parallel series of any pre-existing series of adult shells, except among Nautiloidea ; consequently the angustisellate peculiarities of the ventral saddles and deep lateral lobes characteristic of the latisellate and angustisellate em- bryos of the Devonian and Trias were not due to inheritance from primitive adult radicals, but were later modifications originating in the cecosiphonula from close coiling. They were correlative with the earlier or accelerated development of the depressed whorl, and the quicker growth in bulk of the whorl. Similar tendencies have been observed repeatedly in different progres- sive series of Nautiloidea. Thus, wherever we have been able to trace the series of species from a straight, or loose-coiled, to a close-coiled nautilian form, this as a rule has more complicated sutures. The universal result of such progres- sive specialization among the adult forms of Nautiloids is closer coiling, due to quicker growth in bulk of the whorl, and is accompanied also by the evolution of a larger ventral saddle. It is not surprising that similar mechanical results should follow in the septa of the embryos of Ammonoidea, when similar changes in the mode of growth occurred through the accelerated inheritance of the depressed anarcestian radical whorl, and closer coiling in the cecosiphonula. Branco has observed the shortening of the larval stages in the Latisellati as compared with the Asellati, and the still greater acceleration of development occurring in the Angustisellati, and the correlation of these with the general pro- gress in complication of the sutures of the adults of the same divisions in time. This confirms our previously published opinions of the relation of embryos and adults, and also agrees with those here published regarding the inheritance of the primary, radical, smooth form in the depressed embryos of Latisellati and Angustisellati, and the correlative evolution of the sutures and coiling. The microsiphonula appeared in the Ammonoidea with the second septum, in what is morphologically the second air-chamber when compared with Nauti- lus, though actually the first existing in the apex of the true conch. This microsiphonula is also an accelerated form, since the siphon becomes very rapidly or even abruptly attenuated. The collar or distinctive organ of the siphon among the normal Ammonoids was not formed until later, though the precise period 3 18 GENESIS OF THE ARIETIDA. has not been ascertained in any one form, so far as we know. The microsi- phonula occurred, as might have been expected, earlier than the true goniatitic stages, or goniatitinula, in those species which had the nautiloidean stage with ventral saddle also prolonged into the second septum, as in the Asellati figured by Sandberger, and Gon. atratus figured by Branco. The goniatitinula became distinguishable when the first ventral lobe appeared. This was undivided, as in the lower Anarcestes and in the Magnosellaridz among Goniatitine. This stage is prolonged through one or more septa in the higher Goniatitine, and also in the Lytoceratinze and Ammonitine, and the whorl also at this time strikes one as similar to Anarcestes, or depressed semilunar in section, as stated above, and in these the goniatitinula is completed. The duration of the nzepionic period can in a general way be described as coincident in extent with the duration of the smooth shell, which is always found at the centre of the umbilicus, however much the shell may be subse- quently ribbed and ornamented. This period would of course include many more transformations than the goniatitinula, especially among the higher and later occurring species of the Mesozoic. Haeckel designated all of the progressive stages which succeeded the true ovarian stages and included the nepionic and nealogic stages, and their structural relations, under the term Metamorphology.t This term is, however, somewhat indefinite and artificial when limited in this way, since the ovarian stages are necessarily of very different duration in distinct groups, and cannot be considered as the natural limit of the embryologic period. We should, as above stated, be disposed to think that some such limit as here proposed would be nearer to the true one, namely, to consider the typembryos as the last of the true embryologic stages. This nomenclature would enable an author to give an approximate idea of the stage at which the metamorphologic stages began in any type. Thus, they would have begun in Nautiloids with the asi- phonula, and in the absence of this among Ammonoids with the czecosiphonula. In the absence of this last, if it is absent among the lower Sepioidea, the meta- morphologic stages, according to the same rule, would begin with the first stage immediately succeeding the protoconchial stage. Whenever this last is absent, as it certainly is among the highest of the Sepioidea having meroblastic ova, then its equivalent stage, which represents what is left of the veliger, should be taken as the last of the embryologic stages. As has been noted above, the nepionic period is always smooth, and is visible at the centre of the umbilicus in most discoidal shells, and the demar- cation is therefore visible between this and the nealogic period; but, as can be observed on most specimens, an attempt to separate the characters of the latter from the characters of adults is attended by greater difficulties. It is, however, essential to distinguish the category of ephebolic or adult characters from the nealogic, because in each form of any series there are usually found certain novel characters, which appear for the first time in that particular series. These make their first appearance almost invariably during the ephebolic period. 1 Morph. d. Organismen, IT. p. 22. NOMENCLATURE OF STAGES OF GROWTH AND DECLINE. 19 The ephebolic characters are as a rule inherited or homogenous within the special series in which they originated, but are not transmitted from one series to another except through the medium of the nealogic stages of what we have called the tertiary radicals} and they are not, so far as we know, ever concen- trated in the earliest larval or nepionic stages; they occur too late in the history of types. We classify in the nealogic and ephebolic stages such characters as follows: the sharply defined ridge-like pil and tubercles, the channels with their lateral ridges, and keels, and especially the hollow keel, the highly developed rostrum of the higher suborders, especially Ammonitine, the lateral lappets of the aper- tures, and the branching marginal lobes and saddles of the sutures of suborders above Goniatitine. Speaking in a general way, we should include in these categories those progressive characters which appear late in the life of the shell among the higher suborders, and at the acme of their development in time, which are not found in the stock of discoidal radical forms. When the shell began to assume the ribs or pilex, as we prefer to call them, the nealogic period may be said in a general way to have been entered upon. It has been found that these stages of growth indicated genetic relationship with radical forms, which were not infrequently merely different genera or species within the limits of the same family, and often occurred on the same or only slightly different horizons. The nealogic stages of the higher Ammonoids, Ammoni- tinee and Lytoceratin, have not the constancy and general importance of the nzpionic stages, but are transient in the history of the types, appearing and disappearing in the same limited series of forins. They consist of the less im- portant modifications which first appeared in the adolescent or adult stages at a late period in the history of a type, and were then inherited in the nealogic stages at earlier ages in successive species of the same series, according to the usual action of the law of acceleration. The nealogic category cannot be as definitively separated from the characteristics of adults as from those of the larvee. heir first appearance in adults indicated the establishment of a new species in any given series, since they are invariably differences so far as their predecessors and congeners in the same series are concerned. However much they may represent or reproduce the characters of species in other series, they are essentially differentials as regards the adult stages of ancestral species of the same series. Thus the nealogic characters are as a rule ephebolic, and not nealogic, in origin among the Cephalopoda, and usually become nealogic through inheritance. We shall have frequent occasion farther on to call in the evidence of the ephebolic stages, and to show, as in the Endoceratide, that, as a rule, characteristics originated in this stage of growth, as indeed must have been the case with the cecum and the microsiphon. At the termination of the progressive stages, which ended with the full development of the ephebolic characters, the first stage of decline, or the gera- tologic period, began to make its appearance, and became more and more appar- ent as the specimens advanced in age. It was found that, as has been observed * See, for secondary and tertiary radicals, p. 22 et seq. 20 GENESIS OF THE ARIETIDA. in other animals, especially in man himself, the decline was marked by degra- dation of certain characters, and the number of parts undergoing degeneration was gradually increased, until finally the whole of the body was more or less affected. This period has been frequently described by the author in previous publi- cations, and will be more fully described farther on. It is necessary now only to call attention to the fact, that the geratologic or old-age period can be natu- rally subdivided into two quite distinct stages. The first, or Clinologic stage, included the retrogressive transformations during which the nealogic and ephebolic characters became resorbed one after another, usually in reverse order to the succession in which they were introduced during the progressive stages of growth. The size of the whorl also, sooner or later according to the species, showed retrogression during this period. All of these retrogressive tendencies reached their extreme ex- pression in the last and final stage of the ontogeny of the mdividual. In this stage the spines, pila, and often the keel and channels, when present, were lost, and the size of the whorl was so much reduced ‘in all its diameters that it became more or Jess rounded, whatever the angularity of the whorl during the ephebolic period. This stage we have designated by the term Nostologic, on account of the likeness to its own neepionic period, which was finally acquired by the smooth, almost rounded whorl after the loss of its progressive characters. Geratology, or the study of the relations of these old-age stages, shows, as we shall try to demonstrate farther on, that the clinologic characters can be used to predict the degradational modifications which appeared in any series of orna- mented shells when placed under such unfavorable conditions that their descend- ants became degraded, and series of more and more retrogressive forms were gradually brought into existence. A number of such series have been traced by several authors, and they usually end with a perfectly straight form. This form terminated the phylogeny of the series in a manner comparable to that in which the nostologic stage terminated the ontogeny of the individual. It is usually separated also by a gap from all other species, which has not yet been fully filled by intermediate species. This nostologic adult form, the so-called genus Baculites, is not only comparable in this way and by means of its smooth and compressed cylindrical whorl with the last stage of ontogeny, but it is also a very complete reversion to the aspect of the earliest radicals of its own class, the Orthoceratite and Endoceratite. This nomenclature is similar to that originated and published by Haeckel, and at first sight may appear to many naturalists as identical; but it is really only complementary. It is based upon strictly structural and morphological grounds, whereas Haeckel’s nomenclature’ was entirely physiological. This eminent author regarded the ontogeny of an individual as a cycle divisible into three periods; first, the progressive stages of Anaplasis, or those of progressive evolution; sec- ondly, the stages of fulfilled growth and development, Metaplasis; thirdly, those of decline, Cataplasis. He also appreciated and gave full weight to the general physiological correlations which are traceable between the history of a group and 1 Morphol. d. Organismen, IJ. pp. 18-23. THEORY OF RADICALS AND MORPHOLOGICAL EQUIVALENCE. 21 the life of an individual, and in accordance with these ideas designated the pro- gressive periods of expansion in the phylogenetic history of a group as the Hpacme, the period of greatest expansion in number and variety of species and forms as the Acme, and the periods of decline in numbers of species, etc., as the Paracme. Haeckel used also the term Anaplastology for the physiological relations of the stages of progressive growth and those of the epacme of groups, Metaplas- tology for those of the adult and the acme of groups, and Cataplastology for those of the senile stages and the paracme of groups. These terms seem to cover the same ground as those we have employed, but they were in reality chosen for the purpose of classifying physiological relations. Thus the anaplastic relations of the neepionic and nealogic stages to the phenomena occurring dur- ing the epacme of eroups, the metaplastic relations of the ephebolic stages to the phenomena occurring at the acme of groups, and the cataplastic relations of the geratologic stages to the phenomena occurring during the paracme of groups, are the functional relations of the structural modifications occurring in the ontogeny of individuals to those which are characteristic of the phylogeny of groups. THrory oF RADICALS AND MorpPHoLoGicaAL EQUIVALENCE IN PROGRESSIVE ForRMs. The simpler characters of the sutures in the adults of more ancient forms, as compared with the more modern species of the same series, has been noticed by Wiirtenberger, Zittel, Neumayr, Waagen, and Branco,’ in different groups of Ammonitine. The first is very decided in his statement, that the Ammonitinz he has studied form perpetually diverging series, which spring from certain common ancestral forms. The constant repetition of discoidal and involute forms in series, which are otherwise distinct in respect to their sutures and minor characteristics of develop- ment and shell markings, produces a similarity in the succession of the forms. It is practicable to compare the evolution of discoidal into more involute forms of any one series with a similar genetic procession in any other series. Thus in the General Summary, Plate XIV., we can compare the discoidal forms of Ver. Cony- beari, Fig. 20, with Arn. tardicrescens, Fig. 26, Cor. rotiforme, Fig. 30, and Ast. Tur- nert, Fig. 36, and in the same way the involute forms of As¢. Collenoti with Oayn. oxynotum, Greenoughii, and Lotharingum ; and these comparisons also hold good for Schlot. Boueaulliana, and the terminal forms like Woh. Hinumeric: and Psil. mesogenos, which are also involute. In exceptional series the whorls do not become more involute in the higher species, but are nevertheless modified in those character- istics which usually accompany and correlate with increase of involution. Thus the lateral diameter of the whorl decreases, the sides become more and more 1 Wiirtenberger, Stammesgesch. d. Amm., Darwinistische Schriften, Nr. 5, Leipzig, 1880, p. 91. Zittel, Ueber Phylloceras tatricum, Jahrb. d. k. k. geol. Reichsant., 1869, p. 65, Neumayr, Die Phylloceraten d. Dogger und Malm, Ibid., 1871, pp. 347, 348; also, Zeits. d. deutsch. geol. Gesellsch., 1875, p. 866. Waagen, Die Formenreihe d. Amm. subradiatus, Benecke’s Geognost. paleont. Beitr., II. p. 202. Branco, Paleontogr., XXVI., XXVIII. 22 GENESIS OF THE ARIETIDA. convergent outwardly, and the abdomen narrower, though the shell may still re- main discoidal; ex. Caloceras and Coroniceras, Plate XIV. Fig. 11-16, 28-32. The Ammonoids of the Lias also have a tendency to produce keels, ribs, ete. in addition to the parallel procession of the forms just described. Thus, when we study the parallelisms occurring in different series or genera of the Ammonitinze in the same family or group, we find that equivalent species in different series are due not only to the increasing involution of the whorls, but also to the development of similar structural characteristics. Most palseontologists are not aware of these facts, and therefore are apt to consider species of distinct series as closely allied. It is usual, for example, to classify all the species of the Arietidee having quadrago- nal whorls, deep channels, prominent keels, and well developed pile, as species of the same genus, Arietites,’ whereas they are more closely allied to Psi. planorbe, their radical ancestor, than they are to each other. Errors of this kind are common, and have been still more general. Thus most modern improve- ments in taxonomy in all the branches of the animal kingdom have consisted in doing away with classifications made by the association of representative forms, or, as they are here called, morphological equivalents. The Arietidze sprang from discoidal species of Psiloceras, having smooth shells and phylliform sutures. Other groups occurring later in time are traceable to forms of more advanced structure, so far as the shape and ornaments of the whorl and the sutures are concerned. In every case, however, progressive groups have been traced directly to forms having discoidal shells. The discoidal radicals of different series have been invariably found to be nearly related to each other, and to preceding discoidal radical types, while their descendent species are diver- gent, and essentially distinct. However closely they might have resembled each other as morphological equivalents, they possessed the homogenous differential characteristics of their own genetic series. I have elsewhere noted the facts tending to establish the probable existence of a continuous line or radical stock of types or species.” The paleozoic primary radicals are similar to Anarcestes; the mesozoic or secondary radicals are like Dinarites Mahomedanus, Ceratites Sturt, Gymnites, and Psiloceras; they occur largely in the Trias, and are species with discoidal but rather compressed smooth shells. The tertiary radicals, though discoidal, may be highly ornamented with pile and spines, and have sometimes very broad or coronate whorls; they occur largely in the Jura.2 The primary and secondary radicals, if we follow Haeckel’s nomen- 1 Zittel’s Handbuch d. Pal., I. p. 455. 2 Gen. of Foss. Ceph., Proc. Bost. Soc. Nat. Hist., XXIII., 1883, pp 3823-325. 8 Tirolites and Tropites are acmic or tertiary radicals occurring in the Trias. They are certainly coro- nate forms, with pile, tubercles, and open umbilici. If any one will compare the young of Balatonites or Tropites with the adults of the smooth species of Dinarites and Ceratites as figured by Mojsisovies, he will be able to see that the radical stock is a definable series of forms, with characteristics not only shown in the adults of simpler smooth genera and species, but necessarily repeated in the young of more modified species, like Balatonites, Tropites, etc. It must be remembered, however, that all forms will not have the smooth, compressed secondary radical reproduced in their young; many of them lost this, or had it only very slightly, since it was replaced by the broader-abdomened tuberculated tertiary radical, as in the young of Trachyceras aon. The young of Tropites has a form and sutures similar to those of Glyphioceras diadema of the Carboniferous, and the stock of tertiary radicals may therefore be said to have begun even in the Paleozoic. THEORY OF RADICALS AND MORPHOLOGICAL EQUIVALENCE. 23 clature, are epacmic, and the tertiary are what we should call acmic radicals. Cel. Pettos is an excellent example of an acmic radical in the Jura. It stands morphologically and chronologically at the centre of the affinities of the group of Dactyloide and Stephanoceratide, that is, of the larger part of the odlitic Ammonitine. It is, in its relation to these, and to the characteristics of their nealogic stages of development, an epacmic radical, but with regard to Psilo- ceras, and more ancient secondary radical forms, it is a tertiary or acmic radical. It has a flattened abdomen, very divergent sides, like those of Steph. coronatum, and similar aemic radical forms, and a line of coarse tubercles along the sides. Though altogether distinct from Psiloceras, it is also a perfectly discoidal form. The direct descendants of Pettos, which belong properly to the stephanoceran and allied groups, are also discoidal forms, though the series often have inyolute species, such as Maer. macrocephulum, ete. Tertiary radicals in what we propose to call the Pettos Stock, or Spinifera, according to the evidence of the younger stages and the characteristics of adults, have but one row of large spines in adults, and whorls which are very gibbous or trapezoidal in section, that is, with abdomen broader than dorsum. The whorl may sometimes be smooth, with only one row of lateral spines, but is usually strongly pilated, the pile being single on the sides and as a rule bifurcated only on or near to the abdomen. ‘The sutures have a more or less close resemblance to those of Der. Dudressier’, or Cal. Pettos. The line of descent being broken, we shall, in the imperfect list below, give some forms having two lines of tuber- cles. These, however, have young which, until a late stage, show only one outer line of lateral tubercles, as in the adults of the two species cited above. Sfeph. nodosum of the Lower Odlite is the tertiary radical of that genus, and of Macro- cephalites, Sphzeroceras, Morphoceras, Reineckia, Cadoceras, Quenstedioceras, Aspidoceras, Olcostephanus, and Pachydiscus. All of these genera have some forms which are either closely similar to the radical in the adult stages, or else have young with a nodosum-like stage. Pedfoceras athleta has a similar history, though it is like Dactylioceras in its nealogic stages, it has two lateral rows of large spines, and is similar to Asp. perarmatum in the adult. The huge coronate forms of the Upper Jura, like Olcostephanus Gravesiunus, etc., and the single- spined forms like Aspidoceras Hdwardsiamus, and shells like Asp. perarmatum, Rupellense, etc., with two rows of spines, are obviously in the line of stock forms. Jn fact, one can select from the discoidal shells of these groups a more or less. closely allied series of stock forms, from each of which a separate genus or series of genera arose, until we find in the Cretaceous a new beginning in Hopiies Royerianus and Cornuelianus for the species of that large genus, and of Acantho- ceras, Pulchellia, and possibly Holcodiscus and Costidiscus. The cretaceous group, with nodose keels or lines of tubercles in place of a keel, also belong to the Spinifera, but they form a separate phylum connected, in common with such forms as Acanthoceras mammillare, with the Hoplites series, and their radical is also Royertanus. The radical of Cosmoceras, Cos. Taylori of the Lias, is a species with two rows of spines allied to Deroceras armatum, and the adult characteristics of this species are repeated in the young stages of the 24 GENESIS OF THE ARIETIDA. normal forms of that genus. Wiirtenberger has come to similar conclusions, and has traced a large part of the same genera back to the same origin in the work quoted above. We differ in details, and in the way in which we treat the stem of stock forms, but these differences will probably disappear after further re- searches have been made. His book is full of the evidences of careful work, and we do not feel disposed to offer any criticisms until there is an opportunity to publish our own observations in detail. The young forms of the Spinifera in the later neepionic stage, have a very close resemblance to the young of Tropites before the keel appears, and also an obvious reference to Tirolites of the Trias, and to the more remote and possible ancestor, Glyph. diadema, in the Carboniferous. Per. Defranci is the radical of all of the species of the large genus Peri- sphinctes, and has no tubercles in the adult, but in the young there is a prolonged stage like the adult of discoidal coeloceran species, and in still younger stages a pettos-like stage. This genus embraces a very large number of species which have been traced out by Wiirtenberger, and referred by hin to a species closely allied to the one quoted above in the Lower Qodlite. The absence of tubercles, and the rounded form of the whorl in this group, and the frequent absence of the trapezoidal form and tubercles even in the early stages of many species, show that it is distinct from the Spinifera. We propose to designate it by the term Plicatifera. The tertiary radicals of the keeled groups, the Carinifera, as we propose to call them, have also close structural relations, but are modifications of what.we have called the quadragonal form. Nevertheless, in the young and the adults there is a tendency to reproduce the tertiary radical of the Spinifera. This is to be seen in Wiihner’s figures of Caloceras (Arietites) Coregonense, and that keen observer describes the resemblance of the young just before the keel ap- pears to Cael. Peltos of the Middle Lias. Similar facts can be noted in the young of other forms of the Arietidze, but the keeled stage acquires prepotency in the Arietidx, Their quadragonal, keeled, and channelled forms began in Caloceras, and from this genus sprang the similar tertiary radicals of the later Jura. The radical stock is continued by such species, as follows: Amaltheus Hawskerense, Phymuatoceras enervatum, Hildoceras Walcoti, and Huarpoceras Sowerbyi, which last has a modified quadragonal form until a late stage of growth in some varieties. Oppelia hecticus also has in some varieties a quadragonal form until a late stage, though not so discoidal as most of the preceding. In the Cretaceous, there is Schlonbachia tricarinatus and Westphalicus, which are true stock forms of the Cariniferze.? Haploceras, Desmoceras, Silesites, Pictetia, and the like, have tertiary radicals similar to the typical forms of Lytoceras, and belong therefore to the Lytoceratine. 1 Mojsis. et Neum., Beitr., VI., 1888, pl. xxii. 2 Tt should be noticed in this connection that the characteristics of the so-called pettos-like young of the earlier occurring species of the Carinifera are favorable to Mojsisovics’s view that the Arietidee sprang from Halorites. This genus is closely related to Tropites, and the form and sutures of the young of several species in the Arietidze certainly show affinities for Tropites. On the other hand, as we have maintained above, the affinities and gradations of all the species of the Arietide lead us back into Psiloceras, and the alliance of that genus with Gymnites seems to be closer than with any other in the Trias. THEORY OF RADICALS AND MORPHOLOGICAL EQUIVALENCE. 25 Cal. tortile, Cul. lagueum, and Schiot. eatenata, in the Plicatus Stock of the Arietidee, are more closely allied to one another and to Psil. planorbe than are the morphological equivalents among their descendants to one another. However closely the descendent involute forms may simulate one another, their neepi- onic and nealogic stages are generally distinct, and indicate the series with its peculiar differential characters. Arn. miserabile or semicostatum, and Agas. levigatum, are more nearly related to each other and to Psil. planorbe in the Levis Stock than are any of the descendent morphological equiva- lents. There are several forms closely representative of one another, and ap- parently almost identical, among these morphological equivalents. Thus the adults of Ver. Conybeari are apparently very closely allied to Cor. bisulcatum, and to some forms of Asf. Turnert and Arn. ceras » but all of these are more distinet in their nealogic stages than in the adults. The Arietidse present in this respect a similar picture (Summary Plates) to that of the whole group of the fossil Cephalods. Thus the adults of the earlier and simpler radical species, from which the later and more complicated forms must have been derived, are more closely related in structure than any of their adult descendants. The Cyrtocera- tites, Orthoceratites, Gyroceratites, the Nautilini, and the anarcestian Goniatites of the Silurian, are more nearly related in structure and development, in the simi- larity of the adult sutures, the absence of pile and tubercles, and the mode of growth, than are their direct descendants, the genera of the Nautiloidea and the Ammonoidea in the Carboniferous and Jura. The Nautiloids and the Ammonoids had morphological equivalents, but close parallelism is not constant or frequent, and occurred principally among later forms. We have elsewhere discussed this question, and need only notice well known cases; such as the extraordinary likeness of Clydonautilus to the higher forms of Goniatitine due to its divided ventral lobe, of Centroceras to Agonia- tites, and of Subclymenia to Agoniatites, and also the better known example of the Clymeninz of the Devonian and the Aturia group of the Tertiary. Such cases of morphological equivalence are disposed of by the use of the convenient expression, that these are mere analogies. This expression, however, fills noth- ing but a verbal gap. It neither explains parallelisms, nor the confusion they have occasioned and still occasion in our classifications, nor the constant ten- dency of straight shells to become coiled and of already coiled discoidal shells in progressive series to become involute, to whatever series they may belong, or wherever they may be found, thus producing morphological equivalents in great numbers. The only comparison that represents all these relations to my mind is that of a number of divergent branches united at their bases or radical ends into a common trunk. The branches are composed of groups, which, though distinct, and having differential characteristics, are nevertheless similar in the forms pro- duced, and in the order of procession of these forms. The equivalent forms of the larger branches would be admitted to have origi- nated independently of the direct influence of inheritance. We think that this is also true in the smaller series, since in no case can the similarities of the 4 26 GENESIS OF THE ARIETIDA. equivalents, however close, have been derived or carried across the genetic lines of descent from an equivalent representative species of one branch to that of another. Nor could the similarities of such forms have been derived in any series from the radical species, because involved whorls, keels, channels, ete. did not exist in the discoidal stock forms. Parallel series and equivalent forms, also, occur often in such zodlogical and geological relations that any sequence or descent of one from the other is improbable; as, for example, Aturia of the Tertiary, and Clymeninz of the Devonian; or Centroceras of the Devonian, or Subclymenia of the Carboniferous, and Agoniatites which began in the Silurian. These facts speak with great force for the continuity in descent of the dis- coidal shells, and for the existence of a primitive trunk line of generalized radicals, beginning with the earliest times and lasting into the Jura. The uni- versality of the phenomena leads at first to the supposition that we can account for morphological equivalence of species in different series by some invariable law of growth, such as is evidently the cause of the more exact parallelisms which occur between different individuals of the same species. We might con- sider each species as representing a hereditary grade of structure in the develop- ment of a series, just as any period in the life of an individual would represent a stage of development inherited from some ancestral form. We were led into this error at first, but it is an inadmissible supposition in the light of the facts given above. These show, that the representative forms are absolutely new forms in their respective genera or groups, possessing char- acters not found in the stock or chronological trunk of discoidal radicals, and their resemblances are therefore homoplastic, and not homogenous. There are also many kinds of series among fossil Cephalopoda, and in some of these forms similar to those of the Ammonoids and Nautiloids are not pro- duced, as in the Sepioids and Belemnoids. In these orders entirely new modi- fications accompanied equally complete changes in habits and habitat. The crawling and shell-covered, littoral, radical Orthoceras has in these orders be- come changed into a swimming and predatory mollusk, the shell having become internal. It seems evident in these cases, that the forces of the surroundings and new habits deflected the Sepioids and Belemnoids from the more normal course taken by the Nautiloids and Ammonoids, and thus made the repetition of form or equivalence in the shells impossible, except very rarely, and then only in a very limited sense. Such, for example, are the similarities which exist between the internal shell of Spirula and the external shell of Lituites, or between the pseudo shell of the female Argonauta,’ and the true external shell of one of the compressed Ammonitine, like Cosmoceras or Hoplites. The disappearance of the siphon in the Sepioids, and the naked young of the existing forms of this order, show that too much weight can hardly be given to the modifying and eventually controlling influence of changes of habit, or, what is the same thing, the effects of the surroundings in any new habitat, whether 1 See Evolution of Cephalopoda, Science, III., No. 52, 53, 1854; Foss. Ceph. of Mus. Comp. Zool., Bul- letin, I., No. 1; Proce. Am. Ass. Adv. Sci., XXIII, 1883, p. 341. THEORY OF RADICALS AND MORPHOLOGICAL EQUIVALENCE. 27 sought by the animal or forced upon it by geologic changes. Professor Cope, in his masterly work on the “ Origin of the Fittest,’ and in pamphlets previously published, described “homologous” and ‘heterologous” series equivalent to what we have called homoplastic and homogenous series, and gives numerous instances from all departments of the animal kingdom of exact and inexact parallelisms sustaining the position taken above. This eminent author discusses at length the location of growth force due to use or habits, and shows this to be an efficient cause of modification, thus bringing out clearly and demonstrating a new law of variation. His opinions with regard to ‘mimetic analogy” in external and internal characters differs only in so far as we have preferred to use the term morphological equivalence, because we thought it expressed more exactly the phenomena of homoplasy. He says (p. 96), “I believe such coin- cidences express merely the developmental type common to many heterologous (homoplastic) series of a given zodlogical region.’ With regard to the effects of habit, we should also refer to Cope’s remarks (p. 198), and examples with which he explains the origin of generic characters in the ossification of the cranial walls in the Batrachia, and the origin of horns among Ruminants, as due to habits of defence, concluding (p. 200) that the use of the angles of the parts in question (the head) would result in a normal exostosis of a simple kind in the frogs, or as horn cores in the Ruminantia. Waagen, in his “ Jurassic Cephalopoda of Kutch,” * has made a valuable contribution to the facts in tracing several par- allel series of Lytoceratinee in India and Central Europe. ‘The most important facts which result from the investigations explained in the present volume are these two: first, that in Kachh, in the same manner as in Europe, developmental series exist, which are in part identical with the European ones; and second, that the succession of the identical species in time during the jurassic period in Kachh has been governed by exactly the same laws as have been observed in Europe.” “or facts (parallel series*) like those mentioned, which would be augmented by a good many instances if other groups of Ammonites were as well known as Phylloceras, the only explanation is, that the changes of form in the organic world were dependent upon laws which were innate in them and had not to rely exclusively on outer circumstances. The latter factors, as struggle for existence, geoyraphical separation, etc., certainly influenced the production of forms greatly ; but the fundamental law upon which these influences acted very likely was not the law of variation, as stated by Darwin, but the law of develop- ment, or the tendency of the organisms to produce an offspring varying in a cer- tam well defined direction. If this law be true, the time will come when we shall be able to indicate @ priori, with tolerable certainty, what species a given form can or might produce.” 1 Origin of Genera, Proc. Acad. Nat. Science, 1863; Methods of Creation, Ibid., 1871, p. 229; and Origin of the Fittest, p. 95 et seq. ? Paleontol. Ind., Juras. Fau. of Kutch, I., Ceph., pp. 242, 243. § Waagen’s parallel series end in the evolution of identical forms or species from or through different species. We have never met an example of this kind which did not admit of explanation as the result of migration. Waagen’s remarks, however, apply to parallel series in general, whether the forms ultimately eyolyed are the same, or merely resemble one another more or less closely. 28 GENESIS OF THE ARIETIDA. We reproduce this conclusion in full, though, as may be seen by reading the preceding pages, we differ essentially as to the causes that produced parallelism between different series in the same or different localities. Nevertheless, Waagen agrees with us in rejecting the doctrine of natural selection as a fundamental cause of parallelism, and has also stated in 1875, from independent observations, the possibility of dog what we have been putting in practice ever since 1866; namely, predicting what sort of species would be found as descendants of certain given forms, and then subsequently finding them. This experience has also been shared by Professor Cope, who makes similar statements of his own obser- vations among fossil and recent Batrachians and Reptiles. The method pursued by both of us differs from that ordinarily used by naturalists in predicting the existence of new forms, in that it relies upon the action of the law of accelera- tion, and the constant recurrence of similar forms in different series of the same stock, or, as we have explained above, upon the law of morphological equivalence.’ TuErory or RapicaAts AND Morpno.ogicaAL EQuivALENCE IN RETROGRESSIVE Forms. There are certain species among complicated acmic forms which became the ancestors of uncoiled degenerate series, that can be properly termed nostologic forms on account of their complete reversion to the uncoiled forms of the radical groups among Nautiloids. These were not confined to any special class of forms, though more frequent among the higher than among the trunk stock of radical forms. They are what we have called geratologous radicals. Thus Lobites of the Trias must have sprung from some geratologous radical among the Goniati- tine; and Hauer’s Cochloceras with its turrillites-like whorls, and the straight Rhabdoceras, both have sutures which indicate derivation from some genus like Helictites or Choristoceras among Ceratitinz of the Trias, ribbed shells with very simple sutures.’ Choristoceras, also, had discoidal species in the Rheetic beds. We treat these forms as probably degradational, because of their simpler ornamentation and sutures, and also because the similar uncoiled shells among Gasteropoda and among Ammonitine may be followed until they grade into closely coiled and more complicated shells, from which they probably arose.’ The geratologous forms have a most important bearing on the conclusions reached in this essay. They terminate the geologic history of their suborders, just as the Turrillites and Baculites, and others, appear as the final forms of Ammonitine. They were also coextensive with the existence of the cephalopod type, and were evidently liable to be evolved at any time in their history, and to increase in 1 The law of acceleration and of morphological equivalence has been stated in the Preface, pp. iv. and v. ? These lines were written before Zittel’s superb work, ‘‘ Handbuch der Paleontologie,” had appeared, in which (p. 431) he associates these forms in exactly the same order. Although his text does not allude to the genesis of the forms, his mode of arrangement shows that he probably had the same idea in mind. 5 Parallelisms of Individuals and Order among Tetrabranchiate Mollusks, Mem. Bost. Soc. Nat. Hist., I., 1866-67, and Proceedings of same, I., 1866, p. 302. Genetic Relations of Stephanoceras, Proc. Bost. Soc. Nat. Hist., 1876, XVIII. p. 380. Also Genesis of Tertiary Species of Planorbis at Steinheim, p. 8. THEORY OF RADICALS AND MORPHOLOGICAL EQUIVALENCE. 29 numbers whenever conditions became unfavorable to the evolution of normal progressive forms. The degenerative nature of the uncoiled Ammonitine and Lytoceratinze of the Cretaceous has been very generally recognized. They were regarded as diseased forms by Von Buch and Quenstedt, and Neumayr’s dis- covery of the prevalence of simpler sutures even in the normal forms of the Cre- taceous has completed this wonderful picture of wholesale degradation. It can be confidently stated, that the well known cretaceous genera of uncoiled shells, Crioceras, Ancyloceras, Ptychoceras, Hamites, and Baculites, are the morpho- logical equivalents of similar forms occurring earlier in the Jura, but that they are not their lineal descendants. The series of Cosmoceras (Amm.) bifurcatum worked out by Quenstedt,’ and studied also in detail by the author, had shells which became gradually uncoiled. Quenstedt named the uncoiled forms Ham- ites, but has correctly traced them to the coarsely tuberculated species Cos. bifurcatum. There is also a finely tuberculated specimen, bacu/atus, with a broader abdomen, which does not otherwise differ from defurcatum. To this last he is disposed with good reason to refer an arcuate and a straight baculites- like shell. This same tendency is observable among the shells of the Planorbidee at Steinheim.’ Among living shells of a closely allied, if not identical, species of Planorbis at Magnon,’ similar but exaggerated and evidently diseased forms oceur, and the physical conditions are such that we can attribute the tendency to the unfavorable and abnormal nature of the surroundings. We have previously pointed out, that such uncoiled shells could not have had the same habits as closely coiled ones. The appearance of a rostrum in the Ammonitinz indicates that they had become exclusively crawling ani- mals, in consequence of the disappearance of the ambulatory pipe or hyponome. In the shells of uncoiled Ammonitinze the rostrum though smaller is still present. Scaphitoid, ancyloceratoid, hamitoid, and ptychoceratoid shells, to whatever gen- era they may be eventually referred, have one peculiarity in common, the liv- ing chamber is bent backwards, foumnini a shepherd's crook. The absence of the hyponome and the presence of the rostrum in these forms show that they could not have been swimmers, like the modern Nautilus with its large hyponome and corresponding sinus in the aperture and in the strixs of growth along the outer (ventral) side of the whorls. The shepherd’s crook added to the rostrum in the living chambers of the shells mentioned above indicates not only a wide departure in habits from the close-coiled Nautiloids, but also from the close-coiled Ammo- nitinge, since such creatures could not have crawled with facility. They must have been stationary, either hanging among the branches of plants and feeding upon them, or living with their lower portions buried in the ground and cleaning the surrounding surfaces for their food. Other suppositions might be made, but all hypotheses would involve a wide departure from the habits of their immediate ancestors, and from those of their morphological equivalents, Lituites, Gyroce- ratites, or other uncoiled Nautiloids, none of which have the reversed shepherd’s ? Der Jura, p. 400, plates ly., Ixxii.; Amm. Schwab, Jura, p. 576, plate Ixx. 2 Gen. of Plan. at Steinheim, Summ. Pl. ix. * Ann. Soc, Malacol, Brussels, VI., 1871, Planorbis complanatus (forme scalaire), by M. Lois Piré. 50 GENESIS OF THE ARIETIDA. crook in the living chambers or the rostrum. These cases also illustrate Dohrn’s theory! of change of function, and the effects produced upon organs thereby, which has been of the greatest use in our researches. Semper’s researches and experiments? explain changes in organisms in the same way, as probably caused by changes in the surroundings which have led to the adoption of new habits, and the consequent modification or suppression of already existing organs, and some- times to the building up of entirely new organs or parts. It is interesting to note, that our investigations, though necessarily confined to purely morpho- logical phenomena, have led to theoretical results similar to the conclusions of Dohrn, Semper, and others. We can account for the existence of the parallel series on the basis of the following law of relation to the surroundings: The response or reaction of the forms of different series to the action of the ordinary surroundinys in the same habitat produced progressive morphological equivalence, when the external influences were favorable to growth? The enviroment may assuredly be assumed to have been favorable in the case of the parallel series of normal forms of the Ammonitinze and other chambered shells, whether occurring in India or Europe. The diversity of these causes was very considerable, but it was not of such a nature as to imply a change of habitat, or any fundamental change not favorable to the growth of the shell. The average size of Goniatitinze is considerably below that of the Ceratitine, and these in turn, as well as the Lytoceratinse and Ammonititins of the Trias, are smaller as a rule than the same suborders during the Jura and earlier Cretaceous, The steady increase in size in all the progressive series of the Arietidee culminating in the huge shells of Coroniceras shows this very plainly, as may be seen upon consulting the Summary Plates, and the same is true of Planorbide at Steimheim. When the environment, however, became unfavorable to growth, we find retrogression and retrogressive equivalence. Lobites is a genus of small species; Choristoceras, Cochloceras, and Rhabdoceras are also smaller than most of the Ceratitine. The deformed species of the bifurcatus series are smaller than the normal bifwreatus. All of the scaphitoid shells are notably smaller than their congeners, and though there are many large Crioceratites, Ancyloceratites, and Baculites, there are, so far as we know, no exceptions to the rule in cases which have been traced to close-coiled forms. Retrogression is also exlibited in the decreasing size of the retrogressive forms of Agassiceras, Asteroceras, and Oxynoticeras. In the pathological species with extremely retrogressive forms there is an evident exhaustion of the normal powers of growth and development, and prema- ture senility. This is shown in the uncoiling, destruction of the ornaments, and often also by the retention of nzepionic and nealogic characteristics in adults. The form and sutures of straight shells in the Jura and Cretaceous, for example, 1 Der Ursprung und der Princip des Functionswechsel, Leipzig, 1875. 2 Wachsthum’s Beding. d. Lym. stagnalis, Verhandl. d. Wurzb. phys. med. Gesell. N. F., IV.; also Naturl. Existenzbedin. d. Thiere, Leipzig, 1880; and Animal Life, etc., Appleton’s International Scientific Series, 1881. 8 See also Preface, pp. iv, and y. THEORY OF RADICALS AND MORPHOLOGICAL EQUIVALENCE. Sil differ but little at any age. The four or six lobes of the young are retained throughout life, and have comparatively simple margins. The adult, however, is not similar to a true larval form except in the same sense that an old whorl is similar to its own young, or the toothless gums of an old man are similar to the same parts before the teeth appear. The Baculites are not as a rule strictly tubular whorls, as in the nzepionic stages of other Ammonitine, but are gener- ally more or less compressed in the adults, like the aged whorls of close-coiled shells. The development skipped the normal progressive stages of the proximate close-coiled ancestors, and, like syphilitic children, these shells had no proper adult stages, but assumed senile, degradational characteristics while still young, and are, as we have said above, purely nostologic forms. The law of evolution for geratologous forms seems therefore to be as follows: The response or reaction of the forms of different series to the action of the ordinary surroundings in the same habitat produced retrogressive morphological equivalence, when the external influences were unfavorable to growth. We cannot account for the number of uncoiled Ammonoids in the Upper Cre- taceous, their wide distribution, and the undeniable fact, that they were the members of an order then rapidly nearing extinction, unless we imagine the gen- eral conditions of life during this period to have become unfavorable. ‘The unfavorable causes produced in the forms of the groups as a whole similar modifi- cations to those caused by the unfavorable effects of the local surroundings in Cos. bifurcatum, and other shells in more limited localities during the jurassic period. The bifureatus shells and the uncoiled cretaceous Ammonoids are not isolated individuals, —like the turrillitic deformities of Ammonitine figured by D’Orbigny under the name of Zr. Boblayi, Valdau, and Coynarti, or the planicostan deformities figured in Cor. rotiforme, Plate ILI. Fig. 7-13, and the scalariform Planorbidee of Magnon, or multitudes of similar instances known to every student of these fossils, — but series of varieties, species, and genera. These can only be accounted for as the result of hereditary tendencies acting upon races and species, through successive generations, for periods of time more or less prolonged. The evidence is very strong, that Baculites, Scaphites, etc. of the Cretaceous are not necessarily species of the same genus, but probably always polyphylettic in origin. The Baculites of North America have so close resemblance to those of Europe, that they are usually considered as allied species; but there are indications in the peculiar nodular markings and great size of many species, which lead us to think that they origmated from American stocks. Several species of American Scaphites have common characteristics in the sutures, and in the aspect of the ribs and tubercles, and the abdomen, which suggest affinity with Placenticeras. Meek’s plates of Scaphites, published in his Invertebrate Paleontology,’ exhibit common characteristics so far as the sutures are concerned, especially the large size and length of the first lateral lobes, but he gives no figures of the tuberculated young of Placenticeras placenta, which make this com- parison closer. The Amm. Mudlanus on Plate LVII. Fig. 1, 1 a, from Upper Cre- taceous, Chippeway Point, near Fort Benton, has exactly the form in some 1 U.S. Geol. Survey of Territ., Hayden, IX., plate xxxiy., and Placenticeras on plate xxiii. 52 GENESIS OF THE ARIETIDA. examples; also the sutures, Plate VI. Fig. 9, of the young of Seaph. ventricosus, Plate VI. Fig. 7 b, 8-8 b, of the same locality; the pile, the involution of the whorls, and the sutures are also similar. It differs only in possessing the seaphi- toid living chamber, which is well marked. This group of Scaphites are stouter, and have different sutures from Placenticeras.’ In Europe Stephanoceras refractum, the Anim. refractus of authors, is a true Seaphites, but no one thinks of calling it Scaphites, and it is usually referred to the group of normal Ammonitine, in common with several other distorted forms. In an article on “Genetic Relations of Stephanoceras,’* we discussed the affinities of this and similar distorted forms, trying to show the former existence of a general tendency to imitate the scaphitoid mode of growth in Stephanoceras Gervilii, microstomum, and platystomum. These species rebuilt a living chamber at each arrest of growth, which was eccentric, having a flatter curva- ture, and being smaller than the included whorl. This living chamber was also resorbed at each period of renewed growth, as in Scaphites. The well known form Amm. vertebralis Sow., of the Upper Jura, was described by Quenstedt as a diseased scaphitoid form, derived from Arun. cordatum, and this conclusion has also been confirmed by my own observations. Slephanoceras bullatum® has a shell which is precisely like typical Scaphites in the form and aspect of the last whorl, but does not depart from the spiral as in Scaphites. It is, in other words, intermediate between Scaphites and the normal closely coiled Ammonitine of the Stephanoceran group. The Geyer,’ and Herbich,’ have also thrown a strong light upon the peculiarities of the faunas of the eastern part of Europe, particularly the basin of the Northeastern Alps. All of these researches, and many others not mentioned, have made still further advances in the classification of the chrono- logical relations of the minuter subdivisions or beds practicable. 1 Roches des Jura, pp. 23, 162, 173, et seq. 2 Die Jura-Format. Eng. Franky. u. d. siidwest]. Deutschl. Wiirtt. Jahresb., XII.— XIV., 1856. 3 Die Cephal. a. d. Lias d. norddstl. Alpen, Denksch. Akad. d. Wissensch., Wien, XI. 4 See note 2, page 86. 5 Mojsis. et Neum., Beitr., II. - VI. 6 Ceph. heirl. Schich, Abh. k. k. geol. Reichsans., XII. 7 Das Széklerland, Mitt. Jahrb. d. k. ungar. Anst., V., Pt. IT. 86 GENESIS OF THE ARIETIDA. The principles of geographic distribution first announced by Marcou’ have been carried further by Neumayr,’? who has defined the homozoic bands of life in the faunas of what he has denoniinated the Mediterranean, Central European, and Russian provinces. Neumayr, in his article “Ueber climatische Zonen der Jura und Kreide- zeit,’® describes the boundary between the Mediterranean and the Central Euro- pean provinces. This line, as far as traced by him, begins at the east between the Donetz and the Crimea, at about 47° north latitude, and runs thence to the easterly end of the Carpathians; thence, north-northwest to the neighborhood of Krakau ; thence, southwest towards Vienna, and south of Briinn ; thence, west- erly to the neighborhood of Lake Constance ; thence, west-southwest, and later southwest through southeastern France; thence, across the Gulf of Lyons to Spain, and across that country and Portugal to between 38° and 39° north lati- tude on the Atlantic. This author regards the Mediterranean province south of this line, and the Central European province north of it, as respectively parts of two homozoic bands, which encircled the earth during the jurassic period. The Central European province was defined by Neumayr, in a general way, as Including the British Islands, France, Germany, Bohemia, Moravia, and Poland, north of the line described above, and perhaps the Dobrudscha region. The Jura north of these countries was included in his Russian province, which con- tained Central Russia, Petschora Land, Spitzbergen, Greenland, and perhaps Vancouver's Island in North America. Neumayr quotes the works of various authors upon the fossils found in South America, and concludes that the Jura in Bolivia, Chili, the Argentine Republic, Columbia, and in Central America is probably Mediterranean. He thinks also that the few fossils found in the United states indicate the presence of a Central European fauna. Waagen, in his “ Fauna of Kutch,” shows that India is a distinct basin, con- taining forms of the Upper Jura found in the provinces of the Mediterranean and Central Europe, besides numerous peculiar species. Stemmann is of the same opinion with regard to the fauna of the Upper Jura which is found near Caracoles in Bolivia. We have examined a number of the latter collected by Alexander Agassiz at this locality, also several species collected by him at the pass of Tilibichi in Peru, as well as those mentioned in the chapter “ Descriptions of Genera and Species ” of this work, and have read Gottsche’s “ Paleontology of the Argentine Republic.” These and other sources of information show, we think, the same history as in India; namely, that this region may be advantageously separated as the South American province on account of the number of peculiar species it_ contains. There are, over and above these, also a number of forms identical with those of Central Europe and the Mediterranean. We have also seen the fossils of the Upper Jura, found in California, through the kindness of Prof. Joseph 1 Roches des Jura, pp. 74-91, 280, et seq. 2 Ueber Juraprov. Verh. k. k. geol. Reichsans., 1871, p. 54; Ueber unverm. auftret. Cephal., Jahrb. geol. Reichsans., XXVIII., 1878; and Jurastud., Ibid., II., 1871, p. 524. 8 Denksch. Akad. Wien, 1883, XLVII., and also Geog. Verbreit. d. Jurafor., Ibid., l.., 1885. REMARKS. 87 Leconte, and have collected some forms in California and other localities in the United States. There are also.a few species in the collections at San Francisco, but these and the fossils collected at Vancouver’s Island, and described by Mr. Whiteaves of the Canadian Survey, which we have also seen at Ottawa, show a mixture of the species of the European province besides a number of peculiar forms. Though not disposed to give any final opinion at present, the facts jus- tify the suggestion that the North American assemblage of species has a distinct facies of its own, and ought to be separated at least provisionally from the South American and all European faunas as the province of North America. The collections so far made in the Jura show that there is a prevalence of the Arietidee in the Lower Lias, and of the species of Perisphinctes in the Upper Jura of the South American province, whereas these are less abundant in North America. Whiteaves also shows a mixture of the species of the Cretaceous with those of the Jura at Vancouver’s Island, which, together with the peculiar species found there, suggests a distinct basin for that locality as compared with the Jura farther to the south and east in the United States. The fossils so far found in the district of Atacamas, and at localities in the Argentine Republic, show that a provisional separation should be made between this region and that of northern Peru, and that two basins at least, if not more, exist in the Jura of the South American province. Both the physical features of the distribution of the deposits and the faunas appear, therefore, to make it doubtful whether the terms Mediterranean, Cen- tral Europe, and Russia can be assumed as appropriate names for the homozoic bands of the jurassic period in America. It would be preferable to adopt for these bands the nomenclature of Marcou. Thus, the Bande! Homozoique Cen- trale of Marcou would become the Tropical Homozoic Band; the Bande Homo- zoique Neutrale du Nord of Marcou would become the Temperate Homozoic Band; and the Bande Homozoique Polaire du Nord of Marcou would become the Polar Homozoic Band. These bands could then be subdivided into provinces and basins according to the faunas, and the real facts of the distribution of forms more clearly shown than by using the names of European regions for that purpose. Waagen, in his article “ Ueber die Zone des Amm. Sowerbyi,” has traced in a general way, following out simply the physical features of the distribution of the Jura, the following basins: I. South German Basin, consisting of Suabia, Fran- conia in Bohemia, and southeastern Baden “und des Randen.” II. Helvetic Basin, including Switzerland, departments of Doubs, Jura, and Ain, also Rhone, Saone et Loire, Cote d’Or, Haute Sadne, Haut Rhin, and Bas Rhin, and the neigh- boring deposits in the south of Baden. III. Mediterranean Basin, including the departments of Lozére, Aveyron, Hérault, Gard, Ardéche, Drdme, Basses Alpes, Var, and Bouches du Rhone, and suggests an Italian basin for the Southern Alps. TV. Pyrenean Basin, including the departments of Lot, Charente, Charente Inférieure, and perhaps Deux Savres. V. Parisian Basin, including the depart- ? The use of the word zone instead of band is likely to lead to confusion, on account of its employment in geology for the synchronous faunas of the same beds, and we think it ought to be avoided. oe) (o/2) GENESIS OF THE ARIETIDA. ments to the north, to which Waagen adds Dorsetshire and Wiltshire in southern England. VI. North English Basin, from Gloucestershire to Yorkshire inclusive. VII. North German Basin, including Hanover, Brunswick, and the neighbor- hood of Magdebure. We have not needed to use these divisions precisely as laid down by Waagen, but it is interesting to remark that they accord more or less completely with the observations on the faunas here recorded. Our principal interest has been, of course, in the central portion of each basin, and not in the more deficient records of outlying localities. The South German basin is as it has been given by Waagen. His Helvetic basin appears to be a natural division, with the exception of the departments of Saone et Loire, the Cote d’Or, and the Rhone. The Cote d’Or has appeared to us to be the centre of a different basin, which extended in- definitely through the departments to the westward, and also to the south until it met the fauna in the valley of the Rhone described by Dumortier. Whether such closely contiguous faunas as that of this valley and the Cote d’Or ought to be designated by distinct names we cannot pretend to decide, but that they differ materially from the point of view of the evolution of their faunas seems to us highly probable. The faunas of northeastern France and Luxemburg, though perhaps in a dis- tinct basin from those of Westphalia, Hanover, etc., which are properly included in the North German Basin, are all similar in so far as they contain similar resid- ual faunas. The basin of the Rhone includes the departments mentioned as in the Mediterranean basin by Waagen, with the exclusion of the southeastern part of the department of Var, which, as shown by Dieulefait, belongs to the Italian basin. We have not been able to study any collections from Wiltshire, but the Dorsetshire fossils of the Lower Lias, though certainly presenting a very distinct facies and association cf forms from those of Waagen’s North English basin, have not seemed to require separation into a different basin. The fossils do not resem- ble those of any other fauna so closely as they do that of the rocks in the rest of Envland to the northeast, and, though it may be natural to make this separa- tion, we have not required it for the immediate purposes of this memoir, and have consequently spoken of the entire region as the English basin. The Lias in territories to the north, like Scotland and Sweden, is deficient in Ammonitine, and Judd? remarks upon the estuarine character of the deposits. At Dompau and Déshult in northwestern Sweden a few poorly preserved fossils show the presence of the bucklandian fauna. It is possible that these deposits may have a yet undiscovered fauna of Ammonitinee distinct from more southern localities ; but so far as one can see, the forms of the Swedish basin are not dis- tinct from those of the faunas of North Germany. Neumayr has already traced in a general way the origin of the fauna of Central Europe to the Mediterranean province, and we think a still further advance has been made practicable by the methods of constructing genetic series as advocated in this monograph, and the discovery of definable cycles in the genesis of forms. Though our conclusions have been reached under the dis- 1 Quart. Journ. Geol. Soc. London, 1873, X XIX. p. 98. PSILOCERAS AND CALOCERAS. 89 advantages attending residence at a distance from the fields of research, the results have appeared to be sufficiently novel and suggestive to warrant publica- tion. The results reached have been just what one might have anticipated from @ priori reasoning upon the basis of the theory of evolution and monogenesis, but nevertheless have not been admitted without much hesitation, because of the author's natural feeling that so great exactitude in statement with regard to the relative age of faunas on the same horizon should be distrusted. If our data have led us correctly, there are some basins in the Lower Lias which were capable of evolving new forms. These we have called Aldainic * Basins, because they were centres of origin for new series, and their faunas were what we have called Autochthonous Faunas. Other basins were apparently incapable of giving origin to new forms, or at any rate received all, or almost all, the forms which occupied their territory by migration from the aldainic basins. These we have called Analdainic or Residual Basins, and their faunas Residual or Analdainie Faunas. The beginning of the Arietidse was in the Northeast- ern Alps, and this, being the first autochthonous fauna, was older than all others. Thence South Germany or Suabia was peopled by chorological migration, and then the basin of the Cote d'Or. Thus a Zone of Autochthones, or an aldainic band of basins, was formed running to the westward. North and south of this zone all faunas seem to have been residual faunas. The fauna of the Lower Lias in the basin of the Northeastern Alps was, how- ever, not in the zone of autochthones after the deposition of the Angulatus bed. This zone, just before the deposition of the Lower Bucklandi bed, had become narrowed in its easterly extension, and confined to the faunas of South Germany and the Cote d Or. PsILOCERAS AND CALOCERAS. The discovery by Giimbel? of Psé/. planorboides in the triassic strata of the Bavarian Alps having been confirmed by Winkler® and the shell and sutures figured, there can be no doubt that it is a true Psiloceras. As a result of our researches upon cycles of form, we can, however, unhesitatingly assume that this shell is too involute to be considered a radical of the Arietide. It indicates, if estimated according to the usual history of these cycles, that undiscovered species of disecidal Psiloceratites must have existed in the Trias, as necessary antece- dent or ancestral forms. Two forms have also been cited by Neumayr, in his “ Unterster Lias,” + as Hoc. planorboides and Algoe. form. nov. from the Kossener shales. These are from Wallege, and appear to be the same as those previously cited by Stur® as Amm. cf. longipontinus and later described by Wihner.* Wiih- ner considers them both to be specimens of his Psi/. Rahana, and writes that 1 °A)Oaiva, to make to grow. 2 Ober. Abth. d. Keupers, p. 410. 8 Zeits. deutsch. geol. Gesellsch., 1861, XIII. p. 489, pl. ix. fig. 8. Neumayr also, Unterster Lias, Abhand. geol. Reichsans., VIL., figures this species in the Planorbis bed. 4 Abh. geol. Reichsans., Wien, VII. p. 44. 5 Fuhrer z. d. Excursion. d. deutsch. geol. Gesellsch., Wien, 1877, p. 148. 6 Verhand. geol. Reichsans., 1886, p. 175. 12 dO GENESIS OF THE ARIETIDA. they were probably taken from loose rock not in place, and may have come from a dark gray limestone in the horizon of Psi. caliphyllum or megastoma. They cannot, therefore, be considered forerunners of the psiloceran forms of the Planorbis bed. Neumayr’s and Wiihner’s researches, quoted below in Table VI. and in the chapter on “ Descriptions of Species,’ show that a wonderfully rich fauna of Psiloceratites and Caloceratites existed in the region of the Northeastern Alps ; but, so far as we know, there is nowhere any statement of the appearance in time of the discoidal radical Psi. caliphyllum or planorbe before Caloceras in that prov- ince, as there is in South Germany. The aspect of the fauna is older than that of South Germany ; but though composed of an assemblage of radical forms of Psiloceras, they occur side by side with Cul. Johnstoni and Schiot. catenata (subangu- lare of Wiihner), and are equivalent to the fauna of the Caloceras bed of South Germany, but not to the lowest Planorbis bed. Suess and Mojsisovics, in their table of strata in the mountains of the Osterhornes,' Northeastern Alps, describe a very thick Planorbis horizon, and in the uppermost bed they enumerate Psil. planorbe, supposed to be the English form ; also Psi. Hagenowt and Cal. Johustoni, no fossils having been found in lower beds. Here again it is probably the Caloceras bed, and not the lowest Planorbis bed, which contained the fossils described. In South Germany Psiloceras planorbe, the radical species of the Arietide, is prevalent, as may be seen in collections, and in the works of all the geological writers on this region, especially Quenstedt. Quenstedt notes what he calls the Laqueum layer, and speaks of caloceran forms as having made their first appear- ance somewhat later in the Planorbis horizon than Planorbis itself, and in the “ Ammoniten der Schwabischen Jura” describes and figures a specimen, Planorbis, var. deve, from the Bone-bed, which is placed by most writers in the Rheetic. In the neighborhood of Salin and Besancon, Prof. Jules Marcou has shown that there is a deficiency in the Planorbis horizon, and lately Louis Rollier,’ following in his footsteps, has confirmed these observations. Professor Mar- cou, however, at Boisset, near Salins, found a true Planorbis bed containing the typical species. W. A. Ooster, in the “Catalogue des Cephalopodes des Alpes Suisses,’? enumerates many species; but unluckily the beds are not de- fined. It is, however, evident that the collections in Switzerland which he examined, and the authors he quotes, did not give any data contradictory to Waagen’s conclusions, which we give below. Waagen, in his “ Der Jura Franken, Schwaben und der Schweiz,” says that outside of Suabia, whether going northeast or southwest, one finds nowhere the typical development of the Lower Lias as it exists in Suabia; and it is especially the lowest bed which is apt to be nearly everywhere starved out. This remark and the table given by Waagen are very important, and coincide with the results reached in this chapter. 1 Gebirgesg. d. Osterh., Jahrb. geol. Reichsan., XVIII., 1868, p 195. 2 Form. Jurass. Soc. d’Emulat. Porrentruy, 1883, p. 105. 3 Denksch. schw. Gesellsch. Naturwissen., XVIII., 1861. PSILOCERAS AND CALOCERAS. 91 M. Collenot! mentions Amm. Johnstoni, tortilis, laqueum, and Burgundie as oc- curring in the Planorbis horizon. The collections at Semur show that Planorbis was small, and evidently already losing ground, whereas the fine suites of calo- ceran fossils indicate at least that this series had suffered no loss by migration when compared with the fauna of South Germany. This collection is also arranged to show a bed similar to the Laqueum layer of Quenstedt, called by Collenot the “zone of Amm. Liasicus,’ which contains only caloceran forms, and also Psil. longipontinum. Cal. laqueum is smaller, and more like the German form when found in company with Liasicus? A few dwarfed forms of Psi. planorbe, var. deve, have been found together at Saulieu, and at Beauregard there is a bed with large forms of Oal. Johnstoni and tortile, accompanied by a larger form of Cal. Jaqueum than is usual in South Germany, and a small Ps¢. planorbe, var. leve. The latter is in Boucault’s collection, Museum of Comparative Zoélogy, but not rep- resented at the time of my visit in the collections at the Museum of Semur. The researches of M. F. Cuvier*® are important in this connection. He states that a separable Planorbis bed was found by him on the section of the railway between Arcy-sur-Cure and Guillon, and immediately above this a bed characterized by the presence of Cal. Liasicum. Again, on page 177, he speaks of finding at Gravelles, near Saulieu, a bed containing Psil. planorbe and Cal. laqueum or Bur- gundice, and this agrees with Collenot’s observations. Dumortier! states that Psél. planorbe occurs everywhere in the Planorbis bed of the basin of the Rhone in company with Cal. Johnstoni, though not an abun- dant fossil, and from a fragment in his possession infers that the former may in some cases have reached the great diameter of 220 mm. Quenstedt describes and figures a specimen of Psi. planorbe, var. leve, from Provence,’ which he names Anun. psilonotus provinciulis. Martin® designates the Planorbis bed in the region of the Cote d’Or as the “zone of Amm. Burgundie” (our Cal. laqueum). He considers that the beds of “lumachelle,” the Planorbis horizon, show evidences of having been deposited during a period of violent currents, etc. This is an important fact, since it indicates the littoral character of the deposits. Terquem,’ in the department of Moselle, writes that Ammonites are generally rarer and more often broken than Nautili in the Lower Lias, and enumerates only six species. Chapuis and Dewalque state * that in Luxemburg the Planor- bis zone is not fossiliferous.° 1 Description Géologique de |’ Auxois, p. 209. 2 The remarks of M. Collenot on page 164 are very instructive, and confirm the impressions received from the collections at Semur. 8 Notice Géologique, etc., Bull. Soc. de Semur, ser. 2, No. 3, 1886, pp. 170, 176. 4 Etude Paléontologique du Bassin du Rhone, p. 28, pl. i. 5 Amm. Schwab. Jura, pl. i. fig. 19. 6 Pal. Strat, de l’Infra-Lias de la Cote d’Or, Mém. Soc. Géol. de France, VII. 7 Infra-Lias Luxem., etc., Dept. Moselle, Mém. Soc. Géol. France, V. See also, for similar opinions, Collenot, Deser. Géol. de ’Aux., p. 162, and Dumortier, Etudes Pal. Bass. du Rhone, I. p. 20, IL p. 97. 8 Deser. Foss. Terr. Secon. de Luxembourg. 9 The late researches of Schumacher, Steinmann, and Van Werveke, Erlaut. z. Geol. Uebersichtsk. d. Westl. Deutsch-Lothringen, show that the Planorbis bed containing Psil. planorbe, var. plicatus, is found in the region explored by them, though it is absent in the French part of Lothringen, as stated by Bleicher, Bull. Soc. Géol. de France, ser. 3, XII, 1884, p. 445. In Deutsch-Lothringen it is one meter in thickness. 92 GENESIS OF THE ARIETIDA. In North Germany, according to Schlinbach,’ the Planorbis horizon is pres- ent; but Psil. planorbe is largely, if not entirely, replaced by Cal. Johnstoni, and he designates this layer as the “zoné of Amm. Johnstoni.” Braun” gives similar results for his work in the localities of northwestern Germany; and Emerson, in his essay “ Die Liasmulde von Markoldendorf,” did not find Psiloceras in that basin, though Johnston’ was abundant, and of large size. Romer,’ the first ob- server in North Germany, states that the Lower Lias is less developed in that region than in South Germany, and enumerates only a few species. Schliiter, in his “ Schichten des Teutoburger Waldes bei Altenkirchen,” shows that a thick Planorbis bed occurs in this locality, and Psi. planorbe is abundant, while Cai. Johnston’, which he considers to be identical with planorbis, var. plicata, and Amin. laqueolus, is much less frequent.t He also gives Amm. angulatus as appearing in the upper part of the same bed. There is unfortunately no record of the exact beds in which the fossils occurred, and it is not certain, therefore, whether we are here dealing with the Caloceras bed or a true Planorbis bed. Quenstedt also describes and figures a specimen of Psi. planorbe, var. deve, from Quedlinburg.? The paleozodlogical and geological data, therefore, appear to sustain the conclusion, that Psiloceras and Caloceras, as a rule, arrived later in North Ger- many and Luxemburg, the Céte d’Or, and the Basin of the Rhone, than in South Germany. In England the aspect of the fauna has greater similarity with the Cote @Or and South Germany, than with the North German and Luxemburg basins. The Planorbis zone is well developed, and in the Bristol Museum the South German varieties of Psi. planorbe and the English forms from Cotham® are found side by side. This was also the richest collection in caloceran species which we saw in England, though it was still far behind that at Semur. Rev. J. H. Cross, in his “ Geology of Northwestern Lincolnshire,” claims that no true Planorbis bed occurs, but in place of this a bed containing Amm. angulatus and Johustont, which is probably the Caloceras bed. Wright’s section at Uphill railroad cutting shows the bed containing “angulatus and fragments of Liasicus,” called by him the “ Angulatus bed,” and at Binton, Warwickshire, there is a transition bed con- taining only Zéasicus, included by him in the Planorbis zone.” In his sections of the Planorbis horizon Psi. planorbe occurs earlier than any species of Caloceras at the Uphill railroad cutting; at Binton, Warwickshire ; Street, Somerset; and at Brockeridge and Defford Commons. No mention, however, of Psiloceras in any earlier bed occurs, and its appearance must therefore have been later, as a rule, 1 Ueber Eisen. d. Mittl. Lias, etc., Zeits. d. geol. Gesell., 1863, p. 498; Paleontogr., XIII.; and Die Hannoverische Jura, p. 17. . 2 Der untere Jura in nordwestliche Deutschland, 1871. 3 Verstein. norddeutsch. ool. Geb. 4 Zeits. deutsch. geol. Gesellsch., 1866, XVIII. p. 40. 5 Amm. Schwab. Jura, pl. i. fig. 17. 6 Stoddart, in his “‘ Notes on the Lower Lias of Bristol,’ Geol. Mag., V., 1868, p. 139, shows that Amm. Johnstoni occurs in the section he described earlier than true planorbis, if one can judge from the names he gave to the beds, since no lists of fossils were added. The section given certainly indicates the existence of a Caloceras, rather than a true Planorbis bed. 7 Wright, Lias Amm., pp. 11, 20. WAHNEROCERAS AND SCHLOTHEIMIA. 95 than in South Germany. Tate and Blake! discuss the conditions of the deposi- tion, and arrive at the conclusion, that “it seems probable that no portion of the liassic beds was formed in very deep water, but that even the shales partook of the nature of submerged mud flats.” Psil. Hagenowi occurs in the Northeastern Alps, North Germany, Bohemia, and Switzerland ; and in all these places the true Psi. planorbe, var. leve, is scarce. This form is a degraded modification of planorbe which may have arisen inde- pendently in each locality, and it indicates that this species probably lived under unfavorable conditions in these regions. Caloceras was, however, strongly rep- resented in the same basins. It formed an unbroken procession, so far as Cul. Johustoni was concerned, from the Northeastern Alps to England. The facts, with certain exceptions, of which we shall take note farther on, appear to indicate that Psiloceras was autochthonous in the Northeastern Alps. It probably appeared as a radical or chronologic migrant from the Trias, and gave rise to Caloceras in the Lias. Thence both series may have spread by chorological migration into the basins of South Germany, the Cote d’Or, Switzer- land, North Germany, and England. During these migrations they met with favorable conditions im some localities, and unfavorable conditions in others; hence the inequalities of representation. In both series, however, it is obvious that it was the discoidal species which settled in the new territories to the west of the Mediterranean province. It thus becomes evident that the more hichly specialized and more involute species were probably not the progenitors of any of the derivative series that subsequently arose, — an inference agreeing exactly with all our conclusions with regard to the radical nature of discoidal, as com- pared with involute forms. W #HNEROCERAS AND SCHLOTHEIMIA. The exceptionally rich fauna of the Angulatus (Megastoma) bed given by Wihner’ contains, besides the distinctive Wsehneroceran series, Schlot. angulata, and other forms of the same series, as well as many of the involute Psilo- ceratites and keeled Caloceratites, mentioned above. This assemblage shows undoubtedly that a region so richly populated must have been exceptionally favorable for the evolution of Weehneroceras, and possibly also the autochtho- nous home of Schlotheimia. The announcement by Neumayr® of Waagen’s discovery of a true Schlot. angulata in the Rhetic beds near Parthenkirchen in the Mediterranean province, should be mentioned in this connection. Suess aiid Mojsisovics show that the Angulatus zone is very ‘slightly developed in the Osterhornes mountains, but it contains Psi. longipontinum, Cal. laqueum, Schlot. angulata and Moreana, besides a possible Cor. kridion, identified as similar to that figured by Dumortier in France. This assemblage, therefore, contains the most important of the species found in other regions in the Caloceras bed, as well as in the true Angulatus bed above. 1 Yorkshire Lias, p. 215. 2 Unter Lias, loc. cit., [V., 1886, p. 199. 8 Jahrb. geol. Reichsans., 1878, XXVIII. p. 64, and Abh. geol. Reichsans., VII. p. 44. 94 GENESIS OF THE ARIETIDA. The radical form Schlot. catenata appeared in the Planorbis horizon, according to the collection at Semur, and in this fauna the successive forms of the schlo- theimian series succeed one another without a break in their gradations. Quen- stedt’s work on “ Die Ammoniten des Schwabischen Jura” shows that in South Germany the series may be complete in numbers of forms, and even more re- markable in the size of specimens, and the whole series except Boucaultianus appeared before the termination of the Angulatus fauna. There is also a speci- men referred doubtfully even to the ‘‘ gelbe Sandstein” of the Rheetic beds near Tiibingen, thus carrying the possible origin as far back as in the Northeastern Alps. If, as we have supposed, Weehn. subangulare is found in South Germany, the evidence becomes still stronger that this was the probable centre for the chorological distribution of the group in Central Europe. Schlot. angulata and catenata are very numerous in North Germany; but there is a notable tendency to the production of smaller specimens in the collections we have seen. Schliiter, in his “ Schichten des Teutoburger Waldes bei Alten- becken,’’? states that Amm. angulatus, Moreanus, and Charmassei occur there, but that the latter is never so large as in South Germany. Terquem, in his “‘ Province de Luxembourg et de Hettange,’* mentions Amm. angulatus as occurring in abundance, and of good size, but no species like Char- masse’ or Leigneleti. Chapuis and Dewalque give figures of Schlot. angulata, which show that the species is similar to catenatus in having discoidal whorls and the pile crossing the abdomen. ‘This species is evidently similar to that from Markoldendorf, described by Emerson. Seebach, in his “ Hannoverische Jura,” declares that in North Germany there has so far been found only the Anz. angu- latus (equal to depressus, catenatus, and Moreanus), and denies the existence of Charmasset. Brauns, in his “ Untere Jura im nordwestlichen Deutschlands,” cites both angulatus and Charmasset. The whole series, including radical, discoidal, and involute species, appear to have come into the Northeastern Alps basin first, and to have reached in this locality their highest development in discoidal forms. Thence they seem to have spread somewhat later in time into the Angulatus horizon of the South German basin, and migrated still later to the Cote d’Or and England. In the first two basins they reached their highest development in involute forms, — a fact which strengthens the impression that the series must have originated in the Mediterranean province, since the involute forms are the descendants of the discoidal forms. That they arrived in the Cote d’Or later than in South Ger- many is shown by Tables I. and IL. in which we find Charmassei appearing in the Lower Bucklandi zone instead of the Angulatus zone, and by the presence of two new and more highly modified species, D’ Orbigniana and Boucaultiana, which have not been found in South Germany by any collector up to the present day. These views are further sustained by the fact that the English fauna possesses only a slender representation of the group, all the species being rare, and occur- ring at about the same time as in the Cote d’Or, except Boucaulliana, which is 1 Page 42. 2 Mém. de la Société Géol. de France, V. ® Descrip. Foss. Terr. Secon. de Luxembourg. VERMICERAS. : 95 found somewhat earlier in the Upper Bucklandi bed. Neither Schlot. D’ Orbigni- and, nor any of the similar modifications so well exhibited in the collections at Semur and in the Boucault collection of the Museum of Comparative Zodlogy, are represented in this fauna. VERMICERAS. Vermiceras is represented in the Northeastern Alps by Ver. Conybeart, figured by Hauer, and Ver. Mierlatzicum, Geyer, a dwarfed species. Amm. spiratissimum, Hauer, occurs in company with Conybeari in the Lower Bucklandi bed at Enzes- feld; but this is probably a species of Caloceras, similar to Cad. carusense, of the large variety which occurs in the Bucklandi horizon in South Germany. Suess and Mojsisovics do not ‘give any species of this genus as occurring in the Osterhornes mountains, and this is also the case in several other localities where the formations are sufficiently well developed to lead one to expect that the genus would be represented if at all common in this province. Cal. prespira- tissmum, in the Angulatus bed of the Kammerkahr Alps and Adneth, as given by Wihner,' is the only example of a transitional form. Nevertheless the great development of caloceran species in the Mediterranean fauna shows that a com- plete series of transitional forms probably occurred in that province. Giimbel does not mention Vermiceras, in his “‘ Geognostische Beschreibung der Bayerischen Alpen,’ as having been found in the Kammerkahr Alps, unless indeed his Amm. spiratissimum is a true vermiceran form, or similar to Wiihner’s species of prespiralissimum, nor did he find any species of this series in the gray limestones at Gastiitter Grabens. Herbich, in his “Széklerland,’? gives figures and descriptions of Avvzet. mualtcostatus, with both young and adult similar to and probably the same as his Arie¢. Conybeari, all having been found near Also Rakos in the Besanyer mountains. The radical species Ver. spiratissomum made its appearance in South Germany earlier than elsewhere, if we can regard, as seems to us correct in every way, transitional forms like that on Summ. Pl. XI. Fig. 22, though named as belonging to Cul. daquewm, as really closer to Vermi- ceras than to Caloceras. The principal transitions must have taken place in the Caloceras bed of this basin, instead of in the Angulatus bed, as in the Mediter- ranean province. The basins of South Germany and the Cote d’Or are about equivalent in the number of transitional forms, and it is as easy to trace the gradations from Caloceras to Ver. spiratissimum in one locality as in the other. The extraordinary evolution of the series in the Cote d’Or indicates that it must have met with its most favorable home on the bucklandian horizon in this basin. Even on the Tuberculatus horizon several new varieties were evolved, some of which, however, like debiltatus, Rey., must be considered as degradational, and consequently in- dicate the decadence of the genus in this later fauna. According to Dumortier’s figures and descriptions, this genus is represented 1 Mojsis. et Neum., Beitr., V. p. 53. 2 Mittheil. Jahrb. d. k. ungar. Geol. Anstalt., V., Part II. 96 GENESIS OF THE ARIETIDA. by very few forms in the basin of the Rhone, and Ver. spiratissimum appeared first in the Lower Bucklandi bed. In England the number of varieties or forms is not equal to either of the three faunas above mentioned, but the transitional forms are present. Wright, in his “ Lias Ammonites,” gives a section at Red Car, and Amim. Conybeari is cited as occurring in the lowest stratum of the Bucklandi zone. With regard to the Eng- lish fauna, one can see, in spite of the large size and the multitude of speci- mens, that the small number of distinct species and the entire want of autoch- thonous species, or varieties, indicate a purely residual fauna composed of un- modified forms. This basin is north of the zone in which autochthones arose during the Lower Lias, and the basin of the Rhone lies south of this zone, and both are residual faunas. Ver. Conybeart is mentioned by several authors as occurring in North Germany and in Luxemburg, but, so far as we have seen, other forms of this genus have not been cited, and Vermiceras appears to have had but slight development in these basins The facts, so far as now known, are opposed to the inference that this series originated in the Northeastern Alps. On the contrary, it seems more likely that it began in the Caloceras bed of South Germany with a variety of Cal. laqueum, and subsequently appeared as Ver. prespiratissimum in the fauna of the Angulatus zone in the Mediterranean province. The series, however, did not, either at this time or on any subsequent horizon in this province, meet with very favorable conditions for the evolution of new forms. It must be remarked, also, that the variety of Conybeari figured by Hauer and by Herbich has a whorl quite distinct from that which occurs most commonly in Central Europe. It is more like the degenerate variety of Conybeat, which is usually called Bonnard, though apparently of smaller size. ARNIOCERAS. There are quite a number of forms described by various authors as having been found in the Mediterranean province, but they have all been found in hori- zons above the Lower Bucklandi bed. This may be seen by our Table VI, and also in the fact that Suess and Mojsisovics found no species of this genus in the Osterhornes mountains, the beds above the Bucklandi zone being unfossil- iferous, and Paul states, in his article “ Die Nérdliche Arva,”? that only one species of this series was found in the Lias, and this occurred in the beds above the Bucklandi zone. My notes on the collections at Stuttgardt and Tiibingen do not show so rich a fauna as in the Cote d’Or, nor do Quenstedt’s publications indicate so full a development of the series as in that basin. Thus, though the series began in the Angulatus zone, as shown in Fraas’s collection, it did not reach its acme of development in the South German basin. The evolution of Arnioceras in the fauna of the Cote d’Or is exhibited in the Semur collection, and in Boucault’s col- lection of the Museum of Comparative Zéology. The large nuinber of forms in 1 Jahrb. geol. Reichsans., XVIII., 1868, p. 233. “so CORONICERAS. OF the bucklandian horizon, and their early appearance in the Angulatus zone of Cote d’Or, show that this was their most favorable home. We have identified the earliest occurring Semur specimen with Arn. falcaries, but it had some tran- sitional characters allying it with Arn. miserabile and also with Arn. senucostatum. Arnioceras did not appear at all in the Angulatus zone, but in the Bucklandi zone of the Rhone basin, if Dumortier’s work can be considered as authoritative upon this question. This fauna also possesses specimens of much larger size than any found elsewhere, and the series is quite as fully, though perhaps not so richly, represented as in the basin of the Cote d’Or. In England there are certainly fewer species and forms than in South Ger- many or the Cote dOr, and they appear to have been wholly migrants, not possessing the numerous varieties observable in South Germany and at Semur. Only one, or at most two, species of Arnioceras, called either obdiquecostatus of Zeiten, or geometricus after Oppel, appear to have been found in North Germany. Making all due allowances for negative evidence, this appears to indicate a very slight representation of the genus. Schliiter gives, however, a lengthy descrip- tion and figures of Aimm. obliquecostatus as occurring in a bed between the Angu- latus and the Bucklandi zone in the Teutoburger Wald, and his description and figure show that this species may be in reality divisible into several, — one simi- lar to Arn. obtusiforme, one to miserabile or semicostatum, and perhaps another with more marked keel and channels. The forms are confined wholly to this stratum, which may belong either to the Angulatus or the Bucklandi bed. The Luxem- burg fauna was equally poor. This genus, therefore, certainly does not have the aspect, as far as is now known, of having originated in or near the basin of the Northeastern Alps. The evidence is rather in favor of its having arisen from small planorbis-like forms, occurring first either in the Cote d’Or or in the South German basins. At pres- ent the evidence is not determinative, though somewhat in favor of the former basin. The series subsequently migrated to the Mediterranean province, making its first appearance there in the Upper Bucklandi zone. CORONICERAS. In company with the first arnioceran species at Semur is a doubtful form of Cor. kridion, and later in the Scipionis bed a true Cor. kridion is found together with a representative of Cor. rotiforme. Cor. datum also oceurs in company with these, but is the radical of another subseries of this genus. Cor. kridion is cited by Suess and Mojsisovics from the Osterhornes mountains as occurring in the Angulatus zone, and this is not a difficult species to identify. The Coroniceran forms as cited by the same authors in the Bucklandi zone are represented only by Cor. bisulcatum. THauer’s work,’ however, shows that this is probably only a local peculiarity, though the fauna is not so rich as that of either South Germany, France, or England. Dumortier, in his “ Etudes Paléontologiques du Basin du Rhone,” gives Cor. 1 Nordostlichen Alpen, Denk. Akad. Wien, XI. 13 98 GENESIS OF THE ARIETIDA. kridion as occurring in the Angulatus beds, and figures a specimen.’ According to Fraas’s collection, Cor. kridion certamly appeared in South Germany in the Angulatus bed at Méhringen, and Quenstedt declares it to be a rare form in the Bucklandi zone. Coroniceran forms are so numerous in the Bucklandi zone of South Germany and France, that it becomes difficult to determine whether they were more fully evolved in the one or the other of these basins. Wright's tables and lists show that the English fauna was by no means so rich in numbers of species and varieties as either the French or South German ; and this result, notwithstanding the great size and multitude of specimens found in the various localities of that basin, confirms our experience in the study of collections while in England. The works of North German paleontologists show less thinning out of the forms of this series in that direction than in any of the preceding genera. The names Jisulcatus, multicostatus, and the like, occur frequently. This suggests that in bucklandian times the species of the Arietidae had become hardier and more able to survive in the unfavorable localities to the northward, or else the sur- roundings themselves had changed and become more favorable. There is one fact, however, favoring the former as the most probable conclusion. The speci- mens are neither very abundant, nor are they so large, nor so generally dis- tributed in North Germany as in South Germany. The radical of the third subseries of Corniceras, Cor. Sawzeanum, did not appear earlier than the Upper Bucklandi bed in any fauna, not excepting that of the Mediterranean province.? Chapuis and Dewalque show that Cor, Sauzeanum per- sisted in the Luxemburg region, and that Cor. bisulcatum and mudllicostatum were also present; but the number of forms found there are certainly very limited. Schlénbach mentions the usual fauna of the Buckiandi zone in Brunswick, but the species are not so numerous as in South Germany, and no note is made of their abundance. The absence of the Tuberculatus bed, or its unfossiliferous character when present, is noted by Schlénbach, and this indicates a decrease in number of forms as compared with other regions. Brauns in “ Hannoverische Jura,” and Emerson in “ Liasmulde von Markoldendorf,’ show that the coroni- ceran series is represented, but is not remarkable for the number of species, and in most localities, so far as we can learn, the species of this series are not abundant. Shliiter cites Cor. rotiforme and Cor. Gnwendense as occurring in the Bucklandi zone of the Teutoburger Wald, and his descriptions support these results. He alludes to other forms than these species, but does not enumerate them. The poverty of the later beds of the Lower Lias in North Germany, and the constant recurrence of unfossiliferous strata, are characteristics similar to those of the basin of the Northeastern Alps, and these facts indicate that similar un- favorable conditions obtained there. Dumortier’s work enables us to see, also, that in the Rhone basin on the southern side of the Cote d'Or the fauna thinned out. Thus, though Cor. kridion 1 Pl. xviii. fig. 3, 4. 2 See Mojsisovies’s mention of the zone of Amm. Sauzei, Gebirgsgr. d. Osterhornes, p. 199. AGASSICERAS. 99 appeared in the Angulatus zone, the number of species on the bucklandian hori- zon was evidently more limited than in South Germany, Semur, or England. The coroniceran series, therefore, seems to have arisen on the same level in the Mediterranean province, in the South German basin, and probably in the Céte d’Or. The radicals of the subseries, so far as known, do not follow the same law. Cor. datum has not yet been mentioned or described as occurring in any other basin than the Cote d’Or. Cor. Sauzeanum occurs, however, in northwest- ern Germany, according to“Braun, and in the South German, Cote d’Or, and English basins in the Upper Bucklandian bed, though in the basin of the Rhone and Mediterranean province it is not recorded with certainty from any level earlier than the Tuberculatus beds. It is possible that Cor. kridion may have originated in the Northeastern Alps, but Neumayr and Wihner have not yet found this species in their researches among the fossils of the Angulatus zone, and no good figure has been published. The early occurrence and large number of varieties and species in the collections at Stuttgardt and Semur, and the numerous transitional varieties, also show that Cor. kridion found its most favorable home either in the South German or the Cote d’Or basin. The earlier occurrence of the radical of the third subseries, Oor. latum, at Semur, indicates the Céte d’Or to have been the centre of distri- bution for the Bucklandi subseries. The occurrence of Cor. Sauzeanum on the same level in South Germany, Cote d’Or, and England shows, together with the number and variety of the forms subsequently evolved, that the centre of dis- tribution of the Bisulcatus subseries lay in one or the other of these basins. This conclusion accords with the origin and distribution of the parent series, Arnioceras, and derives additional support from this fact. It is evident also, from these facts, that the Mediterranean province must be regarded as having been peopled with migrants from the province of Central Europe, so far as relates to the subseries of this genus, and this makes it more likely that the radical spe- cies of the whole series, Oor. kridion, also arose in this province. So far as known its appearance in the Angulatus horizon of the Northeastern Alps is not sup- ported by the presence of transitional forms, nor by the presence of Arnioceras in the same horizon. The species, if a real /ridion, certainly must be provisionally regarded as a chorologic migrant from the west. AGASSICERAS. Agas. levigatum appeared in the Angulatus zone of the Semur collection, and was represented by numerous specimens in this fauna. It is also attributed to this horizon in the basin of the Rhone by Dumortier, and is well figured by him." In South Germany Agas. levigatum did not appear until the Upper Bucklandi bed. In England and North Germany it appeared associated with planicosta above the Bucklandi horizon. This radical species, therefore, according to our present knowledge, was a migrant in all of these basins, derived probably from the Cote d’Or or the Rhone basin. 1 Etudes Pal., pl. xviii. fig. 5, 6. 100 GENESIS OF THE ARIETIDA. Neumayr! includes what we consider the young of the radical species of Agassiceras in his genus Cymbites, and states that he has not found them in the basin of the Northeastern Alps. Geyer, in his “ Cephalopoden Hierlatz- Schichten bei Hallstadt,” figures and describes under the name of Cymbites a characteristic young form of Agas. devigatum. Hauer’s drawings of Amm. abnormis, in his “ Unsymmetrische Ammoniten der Hierlatz-Schichten,” illustrate typical shells belonging to the compressed variety of the same species, one of them exhibiting the peculiar aperture, and another the gibbous young in the interior. The radical species of the series, therefore, appeared in the Northeastern Alps not earlier than the Upper Bucklandi beds, as in other faunas more or less remote from the Cote d'Or. In the next subseries we find that Agas. Seipionanus and Seipionis are char- acteristic fossils of the Lower Bucklandi bed in the Cote d’Or and Rhone basins, but in South Germany and England they appeared later, together with Agas. stria- ries in the Upper Bucklandi bed, and in North Germany on the same horizon. So far as known, no species of this subseries has been found in the Northeastern Alps. This series, therefore, has the aspect of having first appeared and met with a favorable home in the Cote d’Or or the basin of the Rhone, where its imme- diate radicals are found. ASTEROCERAS. The asteroceran series is represented in the Northeastern Alps, though appar- ently not by many forms. Hauer figures an Ast. obtusum, var. stellare, and we have seen several specimens from this region, but the fauna evidently was not a rich one as compared with those to the westward. According to Suess and Mojsisovics, this species does not occur earlier than the Obtusus bed? in the strata of the Osterhornes mountains, the Adnether-Schichten in which it appears being placed by them above the Tuberculatus bed. According to our classification, however, this curiously mixed fauna may have begun to receive migrants from the west during the time of the lower bucklandian horizon.? The first recorded appearance of Ast. obfusum occurred in the Upper Buck- landi bed of South Germany, and in the similar formation of Luxemburg. M. Collenot, in his table of the forms in the Cote d’Or, quotes only the usual three names, Amm. obtusus, stellaris, and Brooki. Boucault’s collection in the Museum of Comparative Zodlogy shows, however, that probably all the princi- pal forms were present in the basin of the Codte d’Or, and the type of Ast. Col- lenoti was certainly found there. Dumortier shows in his work, that this species was also present in the Rhone basin, but the series of forms in the genus was not otherwise so complete as in the Cote d’Or. The finest series exists in the collection at the Museum of Stuttgardt. This does not have Collenoti, though it 1 Ueber unvermit. auftret. Cephalopodentypen, pp. 63-65. 2 Op. cit., p. 198. 3 The form cited by Wahner, Ariet. stelleformis, an ally of Ast. obtusum, var. quadragonatum, is cited as having been found in the Megasoma or upper part of the Angulatus beds in the Kammerkahr Alps. This is a doubtful matter, since only one specimen exists, and we have therefore allowed the text to stand as written (Wahner, Mojsis. et Neum., Beitr., VI., pl. xxvi, 1888). See also description of this variety, Chapter V. OXYNOTICERAS. 101 does possess Ast, acceleratum, a form found nowhere else except in the Cote d’Or. Quenstedt’s collection at Tiibingen is very fine, and his descriptions and figures indicate a full representation of species, though Col/enoti is not present. Chapuis and Dewalque show that the Luxemburg rocks contain several differ- ent forms of the genus, though they are not so numerous as in South Germany or England. Schlénbach shows that there is an odfusws horizon in North Ger- many containing the usual forms, but only fossiliferous in certain localities, and Brauns publishes similar results in his work. This horizon according to Schltiter does not appear to have been represented in the Teutoburger Wald, unless his Gmuendense bed and the broken beds mentioned on page 48 of his work be con- sidered the equivalent of all the beds between the Angulatus and Raricostatus horizons. The English fauna, according to Wright's “ Lias Ammonites” and the collec- tions examined by me, has all the principal forms, and often very large shells, and there are also, as in the COte d'Or and Rhone basins, representatives of the extreme modification of this genus, Ast. Collenoti. This series had, therefore, a more general development in all the basins we have considered than any of the preceding series, but in spite of this there seems to be a preponderance of forms in favor of England and France. The unusual case of an early appearance of the radical species Ast. obtusum in the Luxem- burg region should have its due weight, but the evidence of an equally early occurrence in the South German basin shows that Ast. obtusum probably made its appearance as an autochthone upon the level of the Upper Bucklandi bed in -the South German basin, and was thence distributed. It is probable that the series subsequently met with more favorable conditions in the Cote d’Or and in England than in any other basin. OXYNOTICERAS. Oxynoticeras oxynotum, the radical species of its peculiar series, appeared in such profusion and with such excessively compressed and involute whorls in the Northeastern Alps, South Germany, the Cote d’Or, and England, that one seems to be dealing with contemporary migrants from some unknown fauna. With regard to this conclusion, however, it may be well to be cautious. The morphological gap is not so great as appears between an adult of a species like Oxyn. orynotum, and Agas. striaries or levigatum. This is indicated clearly by the development of the individual in Ast. obfusum, oxynotum, and Agas. Scipionianium, as we have tried to show in the previous pages and in the descriptions of the genera and species. Oxyn. oaynotum was a species with a highly accelerated development, and in such forms the departure from allied forms took place sud- denly. In consequence of this abbreviated mode of evolution gaps were left in the series which it is difficult to fill. The evidence with regard to the connec- tion of Ast. Oollenoti with Ast. obtusum and the young forms of Oxyn. oxynotun, 1 See young of Oxyn. oxynotum, pl. x. fig. 4, 5, and 14-17, and Summ. Pl. xiui. fig. 9, 10, and compare with Agas. levigatum, pl. viii. fig. 9, 10, and striaries, pl. ix. fig. 14, 15. 102 GENESIS OF THE ARIETIDA. which convinced me of the derivation of that species from Agassiceras, was found in 1873 in the Stuttgardt collection. The specimens of the last named species had been selected by Professor Fraas out of several barrels of specimens of the same species gathered in the same locality. I looked very carefully in all other collections, handling hundreds of specimens, without findmg any duplicates of these forms. Hauer has given, in his “‘ Nordéstlichen Alpen,” ' an involute form, apparently the same as Oxyn. Lymense, and figures of the young, which are, however, in part distinct.2, His Amm. Greenoughii is evidently a member of the same subseries, and identical with the more involute forms of Amm. Guibalianus of Reynés. This sub- series is only sparsely represented in the Northeastern Alps, and its date of appearance is not yet settled. The collection at Semur has this species in the Birchii or Tuberculatus bed. M. Collenot states that this bed in the Cote d’Or basin contains the same spe- cies and is equivalent to the Tuberculatus, Obtusus, Oxynotus, and Raricostatus beds of South Germany and England, and that it is not possible to separate the faunas, as has been done elsewhere. The appearance of the usual forms of Oxyn. oxynotum in great abundance in Southeastern France according to Dumortier, and also of Oxyn. Simpsoni and Lymense, shows that the last named forms may have made their first appearance in France. This is further substantiated by the fact that Oxyn. Lymense, according to Wright, is found more abundantly in the South of England than in the midland counties. The appearance of Oxyn. oxynotum and Lymense in the basin of the Northeastern Alps can be accounted for by cho- rological migration, in the same way that we have accounted for the presence of Asteroceras and others in that basin. The radical species, ozynotum, is cited by Schlinbach® from only one locality in North Germany, and is not mentioned at all by Dr. Brauns in his “Unterer Jura nérdwestlichen Deutschland,” or by Emerson. The second subseries of this genus is completely represented in the fauna of France. Three species only are found in England, none in South Germany, and two in the Northeastern Alps. Apparently none have been found in North Ger- many‘ or Luxemburg. The collections at Semur contain a nearly complete series of forms, and Dumortier has added others occurring in the Rhone basin. The home of the series, therefore, appears to have been in the Cote d’Or or Rhone basin. This is the only series of the Arietidse which overstepped the boundaries of the Lower Lias. Other species have been reported by various authors as occur- ring in the Middle Lias, especially the Jamesoni bed; but these were found asso- 1 Denkschrift. Acad. Wien, XI., pl. xiii. fig. 6, 7. 2 Fig. 6, 7, appear to us to belong to some species of the second or Greenoughi subseries. 8 Bisenst , etc., Zeitsch. deutsch. geolog. Gesellsch., XV., 1863, p. 502. Amm. affinis, however, de- scribed in Paleontogr., XIII. p. 170, pl. xxviii. III. fig. 1, by the same author, is from Middle Lias, Greene, Brunswick, which is very similar to if not identical with Oxyn. oxynotum. We have not cited it in the table, however, since it may prove to be more nearly connected with Oxyn. Oppeli than with oxynotum. 4 Schliiter describes Oxyn. Oppeli of the Middle Lias as occurring at Altenkirchen and Borlinghausen in the Teutoburger Wald, and Schlonbach describes and figures the same from Amberg. FAUNA OF SOUTH GERMANY AND THE COTE D’OR. 103 ciated with Cal. raricostatum, and therefore, according to our classification, are in the Lower Lias. Oxyn. Oppeli and numismale survived in the Middle Lias of Germany, Ozyn. Oppeli alone in the basin of the Rhone, and Oxyn. numusmale alone, in England. FAUNA OF SoutTH GERMANY. — TABLE I. The notable facts brought out by this table are the following. The abundance and concentration of schlotheimian forms in the Angulatus zone, and their early appearance in the Rhetic. The completeness of the Caloceran series in the lower horizons, and the poverty of the faunas existing between the Geometricus or Upper Bucklandi beds and the Raricostatus bed in respect to these series, and also in the vermiceran, arnioceran, and coroniceran series. The asteroceran series reached a high stage of development as regards the number of forms, but is not represented by the extreme modifications noticeable in the basin of the Rhone. The oxynoticeran series is also present, and even passes into the Middle Lias, but has not a full representation of species. FAUNA OF THE COTE D'Or. — Taste II. The Ammonites at Semur were named by M. Reynés, and these names have come into circulation through publication by M. Collenot in his “ Description Géologique de l’Auxois,” and have also been quoted by Zittel and several other authors. Reynés considers many well-marked varieties to be distinct species. This is our principal disagreement with this author, and the following notes, together with the descriptions of species and table, sufficiently explain other differences of opinion. ; Terquem’s figures of Hettangensis’ show a keeled, broad caloceran form with pilz in the young, which belongs somewhere between Cul. daqueum and raricosta- tum. The specimens in the Museum at Semur, identified as Hetlangensis by Reynés, do not agree with these figures. The specimens identified as Dednast belong to several species, and one of these is so exactly like Pirondi, as figured by Reynés in his unpublished plates, that I have quoted this name as a synonym for Johustoni in the table. With regard to the vermiceran series, we traced the relations as follows. Beginning with spiratissimun, the forms appear to grade into Schlenbachi, which represents Conybeari in the Scipionis bed, then into rotator, which is a close ally, if not identical with Amm. caprotinus, D’Orb., and also with the spinous varieties of Conybeart found in Germany. The simpler ribbed forms grade into conybearordes, Rey., which is not very far removed from spiratissimum, thence into true Conybeari, and thence into Breom, which last is a stouter and more robust form. Breont, Rey., exactly agrees with typical Conydear?, and also with German forms of the same name, whereas Conybeari, Rey., 1s equal to our Bonnardi. Bochardi, Rey., has the form and characters of Conybeari during its earlier and adolescent stages, but 1 Pal. Lias. de Luxem., ete., Mém. Géol. Soc. France, V., pl. xiii. fig. 1, a, b. 104 GENESIS OF THE ARIETIDA. has no tubercles. The specimens are large shells, and afford fine examples of the senile stages. Debililatus, Rey., is similar to our lowest transitional forms of Conybearit. It may be a direct descendant of this from earlier times, or, more likely, a degenerate form. This grades into Landriot, Rey. (D’Orb.), which is simply a more compressed form. The occurrence of a form like Arn. falearies in the Angulatus bed at Semur shows that we may anticipate in the future the finding of the radical arnio- ceran forms at this level or earlier. It is also very interesting to note that Arn. Hartmann, of the Birchii or Tuberculatus bed, is a morphological equivalent of raricostatum, being, with the exception of the young, very similar to that species. The more interesting facts shown by this table are as follows. The succession of the forms in the schlotheimian series has remarkable regularity, according very closely with their genetic relations. The caloceran series, though very com- plete in the lower beds, is not so fully represented as in South Germany. Higher up in the Birchii or Tuberculatus bed of Collenot, and probably upon the highest level at about the time the Raricostatus bed of other basins was being deposited, the series had an unusual number of forms. The vermiceran series has a most extraordinary display of varieties, but apparently not quite so full a representa- tion in the lowest beds as in South Germany. Arnioceras is more fully repre- sented in the Bucklandi beds than in any other fauna, and has also many species in the higher beds. The coroniceran series has a similar history, but is not more fully represented than in South Germany. The agassiceran and asteroceran series are also very fully represented, and have the most highly modified species ; the absence of Brooki will therefore probably be supplied at no distant day. The oxynoticeran series has also a complete history, and probably is nearer perfection than is shown in the table, but it nevertheless seems to have had no Middle Lias forms. Fauna oF THE RuonE Basin.— Taste III. The basin of the Rhone is equally important with that of Semur, and we give below a list of Dumortier’s species and their synonyms in the different horizons. Dumortier mentions only Burgundie, and fragments of Johnston’ and planorbis, in what he calls the Planorbis bed. This indicates the possible absence of the lower. beds of this horizon, since this is evidently the fauna of the Caloceras bed. The Angulatus horizon has a fauna less rich in species than that of the Cote d’Or, especially when one considers the large number of localities from which the author’s collections were gathered. ‘The list includes, besides the species given in the table, Amm. bisuleatus, a very doubtful form. It may be a form of Cony- beari, or similar to the peculiar sulcated form described in the note above on page 70, but it is probably not a true Cor. bisuleatum. There are no transitional beds mentioned between this and the bucklandian horizon, and the beds are evidently not so fully presented, either geologically or paleontologically, as in the basin of the Cote d’Or. The list is very meagre as compared with that in the corresponding beds at Semur, but the presence of FAUNA OF THE RHONE BASIN. 105 Scipiomanus indicates that the bucklandian horizon of this basin represents the Lower Bucklandi beds of other basins, Dumortier divided his zone of Amm. oxynotus into four beds, distinguished by their faunas. The “ Davidsoni bed” should have been called Striaries bed, since his Amu. Davidsoni* is identical with Agas. striaries. The list of species does not enable one to synchronize these beds with the Tuberculatus beds of Semur or other basins, nor do they show that it is equivalent to any bed above the Upper Bucklandi beds. The Stellaris bed’ of Dumortier contains, besides the species mentioned in the table, Amm. Locardi, a species of Deroceras, and Ami. Birchi, a form of Microde- roceras; both of these, therefore, belong to a family distinct from the Arietidee. The presence of Birchi, Boucaultiana, and obtusum show that this, and not the so called Davidsoni bed, is the equivalent of the bed immediately above the Upper Bucklandi beds at Semur. This result confirms our opinion that the David- soni bed of Dumortier should be called the Upper Bucklandi bed. Dumortier’s Planicosta bed contains Oluniacensis,? which is identical with As¢. Collenoti; and this seems to settle the geological position of this important species. Amn. seyunus® seems to be an abnormal or diseased Arn. miserabile; Pellati is a young form of Cal. raricostatum ; and armentalis,* if one can trust the aspect of the inner umbilical pile, is a diseased form of Cal. raricostatum. It appears from the figure to be similar to the deformed Am. longidomus ceger of Quenstedt,® and other similar pathological forms, in which the keel and channels have been super- seded during growth by pil crossing the abdomen. Viticola (Plate XX XI. Fig. 9-13) is the same as the Johnstonian variety of Cal. raricostatum ; Hdmundi (Plate XX XIX.) is the equivalent of the young of Cul. nodotianum ; turdecrescens (Plate XX XI. Fig. 3, 4) may be related to Arn. falcaries. The umbilicus, sutures, and general aspect of the last indicate that it is a form of Arnioceras. Oosteri (Plate XXX. Fie. 2-4) is a keeled and channelled form of Arnioceras, with distorted pile. Amin. planicosta, subplamcosta, and Pauli are all varieties of our Der. plamcosta, and belong to a family distinct from the Arietide. The three upper beds of Dumortier are apparently the equivalents of the Birchii or Tuberculatus beds in the table of the Cote d’Or basin. The notable facts brought out by this table are as follows. There is a regularity in the distribution of the schlotheimian series similar to that in the Cote d’Or basin. Caloceras is not so fully represented in the lower beds, and is equally deficient in the Bucklandi zone. It is represented by a full list of species in the highest beds, with the exception of nodotianwm, which is absent. Cal. carusense, however, is more fully represented, and Cal. raricostatum has a greater number of varieties than in any other fauna. The arnioceran series is not so fully represented in the Bucklandi zone, but it is notably richer in forms in the highest beds than im any other fauna. Coroniceras is well represented in the 1 Pl. xxi. fig. 1-4. 2 Pl. xxv. fig. 8-10. § Pl. xxxi. fig. 6-8. 2 Vals xodb-g site 1, Bh 5 Die Amm. d. Schwab. Jura, pl. vi. fig. 3. 14 106 GENESIS OF THE ARIETIDA. lowest beds and in the Bucklandi zone, but is deficient above. Asteroceras is probably more fully represented than is shown in the table, since the extremes of the series have been found, and, the fauna being near to that of the Cote d’Or, there are grounds for anticipating the discovery of intermediate forms. Agassiceras is complete in its lower forms, but Se/piow’s has not yet been found. The oxynoticeran series is not only quite complete, but has also a middle lias representative. As regards Schlotheimia, Caloceras, Vermiceras, Arnioceras, and Asteroceras, this fauna impresses one as containing the most highly modified derivatives, and as being possibly a residual fauna representing an acme of choro- logical migration and varietal modification so far as these genera are concerned. Possibly Oxynoticeras will also have to be included in this category, and then the parallel with the English fauna north of what we have called the zone of the autochthones would be complete. Fauna or ENGLAND.— TABLE IV. In this table the same regularity of succession is found in the schlotheimian series as in the Cote d’Or and Rhone basins. Caloceras is again deficient in the Bucklandi zone, as in the Rhone basin, but is quite fully represented and has an extraordinary new form in the Raricostatus bed, Cal. aplanatum. There is also a curious parallelism with the Rhone fauna in the arnioceran series, which, as in that basin, has the extraordinary form of Arn. Macdonelli of the Raricostatus bed. Besides the general absence of radical species, except of course the generally distributed psiloceran and caloceran radicals, there is in this fauna a very impor- tant fact to be noted, similar to that observed in the fauna of the Rhone. The extreme modifications in the highest formations are very generally present, — more so than in any other fauna. Thus, besides Cul. aplanatum and Arn. _Mae- donelli there are doubtful forms of Cor. disuleatum in the Oxynotus zone. Ast. Collenoti, Ast. denotatum, and the extraordinary series of var. sagillarius of Ast. oblusum, are also present. The Oxynotum subseries is complete, and the second or Guibalianus subseries alone is imperfectly represented. The English fauna is therefore a residual fauna, not only because of the absence of radicals, but because it presents a chronological and biological acme in the evolution of the most highly modified and most recent forms of the different series, thus clearly indicating chronologically and biologically its more recent derivation by chorological migration from the older, though apparently contem- poraneous, faunas of the autochthonous zone. FauNnA OF THE PROVINCE or CENTRAL EuRorE. — TABLE V. This table has already been amply explained, with the exception of certain general facts. The independent origin of the schlotheimian and psiloceran series is in strong contrast with the Northeastern Alps fauna, which as tabulated in Table VI. shows that Psiloceras and Schlotheimia are connected by means of intermediate weehneroceran forms. Schlotheimia and Caloceras are character- wa FAUNA OF THE PROVINCE OF CENTRAL EUROPE. 107 istic of the Planorbis zone; they were immediately succeeded in the Angulatus zone by a full presentation of schlotheimian, caloceran, and vermiceran species, that is, of the entire Plicatus Stock. This stock then entered upon a period of decadence, slight in the-Lower Bucklandi, but more marked in the Upper Buck- landi bed. Arnioceras attained its greatest development in the Upper Bucklandi zone and was more persistent in the higher beds than Coroniceras. This last attained its fullest expansion earlier in the Lower Bucklandi beds, and declined rapidly in the Upper Bucklandi, and disappeared altogether in the Obtusus bed. This decline is shown by the geratologous characteristics of the species in the Upper Bucklandi beds, rather than by a less number of forms. Thus Cor. orbicu- latum, Gmuendense, trigonaltum, and the mudlticostatus variety of bisulcatum, are all degenerate species as compared with the forms of the Lower Bucklandi bed. They have more convergent-sided whorls, and these are usually developed at an earlier age. Agassiceras also reached its acme in the Lower Bucklandi bed, but is more persistent, and has some forms in the higher formations. Asteroceras is the only series which attained its acme in the Obtusus zone, and then declined in the Oxynotus zone. The oxynoticeran series reached its maximum in the Oxynotus zone, and, though surviving the changes which attended migration into middle liassic habitats, became extinct in that formation. The schlotheimian series is a highly modified series, composed of involute derivatives, and ceased to exist in the Obtusus bed, but there are a few dwarfed forms in the Oxynotus bed. Caloceras persisted in the highest beds, whereas its highly modified derivative series, Vermiceras, is shorter lived, and less fully represented in the highest beds. Arnioceras is parallel with Calo- ceras, and is the radical series from which the more highly modified and shorter lived Coroniceras originated. Agassiceras, the radical of the remaining series, persisted from the Angulatus to the Oxynotus bed, whereas the deriva- tive Asteroceras and Oxynoticeras were both shorter lived. These series, even when thus minutely followed out, accord with the law of persistence in radical stocks, as expressed above, on page 26.1 Psiloceras itself is not persistent, and is an apparent exception. It is the last of a long line of paleozoic secondary radicals which survived in the Lower Lias. It can be compared with the upper part of a stem which has reached the point of growth at which it splits into many branches. Psiloceras was in like manner resolved into derivative forms, the arietian radicals Caloceras, Arnioceras, and Agassiceras. We have already noted and discussed the rise and progress of each series: first, the radical stage, or epacme ; second, the acme; third, the final decline, or paracme, caused by the prevalence of geratologous forms. The result of such a serial history, when the series are considered together as one family found within certain specified beds, is shown in this table. There is a precise parallelism between the history of the whole and of any one series. The Planorbis and 1 Tf, as we have inferred above, on page 24, the channelled and keeled species of Caloceras are transi- tional to Hildoceras Walcoti and other radical forms of the Carinifera, this opinion acquires additional strength, since Caloceras would then become the tertiary radical for the whole of the Carinifera of the Jura. 108 GENESIS OF THE ARIETIDA. Angulatus zones contain principally radical species and their immediate deriva- tives. The Bucklandi zone is characterized, with some exceptions occurring only in the Upper Bucklandi bed, by the presence of truly progressive forms. The highest beds, the Obtusus and Oxynotus zones, are almost exclusively the homes of more or less degenerate and geratologous forms. Extraordinary and unforeseen correlations, such as these, between chrono- logical distribution and a biological classification founded upon the life history of the individual, cannot be accidental. We have already shown, in preceding chap- ters, that our classification of series is natural, and capable of verification by means of the cycles which are found to be present in the history of the individual and of the group. ‘The process of verification does not, however, end with this, since approximately exact agreements may be found between the paleozodlogical and geological records wherever both classes of facts exist and have been minutely studied. There is even some evidence that cycles may be traced in the so-called con- temporaneous faunas of the same horizon. Thus, what we have said about the analdainice faunas of England and the basin of the Rhone indicates this possibility. These faunas show an extraordinary evolution of the geratologous forms of the geratologous series; the aldainic basins show, on the contrary, in so far as the Cote d'Or and South Germany are concerned, an extraordinary assemblage of the progressive forms of the Arietidee, whereas the originating or aldainic centre of the family in the Northeastern Alps has a fauna in which the radical series are enormously developed. This would seem almost evidence enough that there are cycles in the chorological migration, as well as in the chronological evolution of forms. 'The whole might be represented as a complex of vortices, in which the result is apt to be a cycle, whether the spiral lines of evolution form small vortices upon the same or nearly the same horizons, or whether the picture is the blending of all these into one great spiral, or a series of more or less parallel and blended spirals ascending through geologic time. FAauNA OF THE PROVINCE OF THE MEDITERRANEAN. — TABLE VI. It was intended to omit this table, as well as those of the North German basin, Italy, Corsica, Spain, ete., the species of which have not yet been fully described and illustrated, since it is not practicable in such researches to accom- plish much unless aided by very full information. Lists of names from which these faunas might have been made up are rarely of much use, since authors differ essentially in the identification of species, and therefore we have not considered it safe to venture upon tabulating them. The publication of Wibner’s and Neumayr’s researches, however, induced the author to attempt to give a tabular view of the Mediterranean province. It has not been found practicable to carry out the system of connecting the forms by lines representing genetic bonds, except in so far as they have been published by the authors named above, and the usual connecting lines have therefore been omitted in series occurring above the Lower Bucklandi bed, and in all the genera of the Levis Stock. FAUNA OF THE PROVINCE OF THE MEDITERRANEAN. 109 The mixed faunas of the Adneth and Hierlatz beds, and of the gray lias limestones and Fleckenmergel, have been described by Giimbel,' by Dionys Stur,” and by Geyer,’ with very interesting remarks upon the similar faunas elsewhere. The first author regards the faunas of the Adneth and Hierlatz limestones as having species representing not only the various faunas of the Lower Lias, but also the faunas of the Middle and even Upper Lias. Oppel con-— sidered the Hierlatz beds as the equivalent of the Obtusus, Oxynotus, and Rari- costatus beds.* Geyer, who has examined this locality in detail, thinks, if it is compared with any single fauna, that we should have to select that of the Oxy- notus bed. He however calls attention to the occurrence of As?. obfuswn and Cal. raricostatum in the same horizon, thus demonstrating the mixed character of the fauna. Stur regards it as possible that the different beds of the Lower Lias may, by further investigation, be defined in the Adneth and Hierlatz beds. This conclusion, however, rests upon theoretical considerations, and not upon actual observations, and this author observes, ‘‘ dass in den Alpen einzelne arten der Lias fauna hoher oder tiefer hinauf und herabreichen als in den ausser- alpinischen Schichten beobachtet wurde, ... und . .. wihrend der Liaszeit innerhalb der Alpengebiets eine weniger strene geschiedene und minder man- nichfaltige Gliederung wirklich vorhanden ist.” ° Both Stur and Giimbel distinguish only three faunas in the Lower Lias of the Kammerkahr Alps: 1. A yellowish limestone with a species similar to Johnstoni. 2. An intensely red limestone with Amm. spiratissimus of Hauer, Li- asicus of Hauer, Huuert, Kridion, Ceras, Bodleyi, Hierlatzicus, Grunowi, bisulcatus, oxy- notus, euceras, Charmassei, acutiangulatus, Doetzkirchnert, Hermanu, Kammerkahrensis, Purtschi, cylindricus, Lipoldi, Foetterh, Petersi, but in which, however, a true Bucklandi bed was not distinguishable according to Giimbel. 3. Above this, thinner layers with Amm. raricostatus, zithus, densinodus, and a form similar to stelluris. Giimbel states that the Adneth or dark red limestones, the Hier- latz, and the gray limestones of Gastatter Grabens are equivalent to one another, and that each contains a mixture of species from Lower, Middle, and Upper Lias. Suess and Mojsisovics® distinguish a Planorbis, an Angulatus, a Bucklandi, a Tuberculatus, and an Obtusus bed in the Osterhornes mountains, but consider the Angulatus bed as the equivalent of the Enzesfeld limestones, and the Obtusus bed as the equivalent of the Adneth limestones. The fauna found by them did not, however, so far as published, appear to justify this conclusion. Wiihner’ gives a clear statement of the facts in his “Heteropischen Differ- enzirung des alpinen Lias.” He quotes Stur® as having distinguished two beds at Enzesfeld, the yellow limestones of the Angulatus zone underlying the true red limestones of the Adneth or Rotiformis horizon. The various localities of the 1 Geogn. Beschreib. d. bayer. Alpen, pp. 428-482. 2 Geol. d. Steirmark. 3 Ceph. Hierlatz-Schichten. 4 Neues Jahrb. 1862, p. 60. 6 Geol. d. Steirmark, p. 433. & Op. cit., p. 195. 7 Verhandl. k. k. geo]. Reichsans., p. 168. 8 See Stur, Lias Hirt. u. Enzesf. Jahrb. geol. Reichs., 1851, pt. 3, pp. 19, 24. 110 GENESIS OF THE ARIETIDA. Mediterranean province are summarized by Wahner in this very satisfactory paper, and one sees that the lowest beds are apt to be well defined, but that after passing through the Angulatus zone definition becomes more difficult, so that even this author, for whom as an acute discriminator of species we have a ereat respect, seems not to have been able to define the separate beds in either the Adneth or the Hierlatz limestones. Herbich makes a valuable contribution to this problem in his §zéklerland, in which he describes several species of the Arietidze, including an Asteroceras like stedlaris of Hauer, equivalent to our obtusum, var. stellare, and Afgoc. Allhii, which appears to be a true Microceras allied to Mier. planicosta, together with a number of species of the Lytoceratide, all occurring in a bed not over three meters thick, and he denies that any distinct beds can be defined.’ Geyer, in the work above quoted, gives a detailed argument for the probable admixture of faunas, and comes to the conclusion that Oppel’s scheme of zones is not applicable to the Northeastern Alps so completely as it is to the formations of Central Europe. Favre, in his “ Terrains Liassiques et Keupe- riens de la Savoie,’ gives a list of localities in which mixtures of different faunas have been announced by various authors, and Geyer adds several other localities. Favre considers that the species in such localities, among which he includes the Northeastern Alps, must have been protected from the geological changes which produced new forms and modifications in other localities, and adds that we must seek the causes of admixture in the continuation of sediments of the same nature, and in the configuration of the surface. His idea was, that the per- sistent species continued to exist in closed basins, where they were secure from the action of the causes that destroyed the faunas to which they originally belonged in other localities. This explanation has a reasonable sound, but it appears to us inadequate. We regard the species quoted as migrants from pre- viously existing faunas, which, having found favorable homes in these localities, became the radicals of new series upon new horizons; or else they were survivors of the geratologous forms of faunas upon the same horizon, which, having found favorable conditions in these new localities, persisted perhaps somewhat longer than the parent series. We have not found adequate evidence of closed areas, except perhaps between the western extension of the Mediterranean province as a whole, and that of Central Europe. The basins of the Lower Lias were evi- dently not, as a rule, so completely closed as to keep out migrants from other basins and provinces, since all the evidence tends to prove the constaney and uninterrupted migration of species throughout the faunas of Central Hurope and the Mediterranean province. Whatever hypothesis is maintained, there seems to be no possible way of accounting for the finding of a species in a truly anachronic position ; that is to say, ina bed which belongs to an earlier horizon than that m which it has been proved to have originated. A specimen of Coroniceras Buckland’ in the Planor- bis bed, or even in the lower part of the Angulatus bed, would introduce great 1 Page 103. FAUNA OF THE PROVINCE OF THE MEDITERRANEAN. 111 confusion into any stratigraphical or genetic classification. We have not yet been able to find any such case! Examples of mixed faunas such as have been quoted above are not so exten- sively mixed as has been claimed. The Hierlatz and Adneth limestones are, . for example, mixtures only of the faunas of the beds above the Angulatus bed; the examples given of so-called psiloceran forms as occurring in them are due to mistakes in identification, since these forms are species or young of species of Arnioceras or Agassiceras, and the species cited as belonging to the Middle and Upper Lias are either radical forms or else morphological equivalents, like all the so-called anachronic forms which we have yet studied. A paper by W. B. Clarke? is very instructive in this connection, since he found in the Rhetic a true Arcestes, showing conclusively how favorable this region must have been for the preservation of ancient forms. He also was able to make out and describe the Planorbis and Angulatus horizons, with a full list of species already described by Wahner and others, and, above this, the Hierlatz horizon. The facts appear also to accord perfectly with the theory of autochthonous faunas. If the Northeastern Alps were the seat of origin for the major portion of the radical forms of Arietidee, we should naturally expect to find in this province the geological and zotlogical relations which are shown in Table VI. ; namely, a clear definition of the lower formations and faunas throughout the Planorbis and Angulatus horizons, and an extraordinary number of radical species and their immediate allies, these also having in the sutures a more ancient or triassic aspect than in Central Europe. An analdainic fauna made up of modified forms arising by migration from other faunas would necessarily be shown either in the admixture of forms above these horizons in case the sediments were similar and continuous, or else in the non-appearance of new radical or progressive forms if the sediments were more varied and more distinctly separable, as in England and in the basin of the Rhone. While the Mediterranean province was an analdainic fauna so far as the Arie- tidae were concerned during the deposition of the upper beds of the Lower Lias, subsequent to the deposition of the Angulatus beds, this was by no means the case with other groups, such as the Lytoceratide. On the contrary, as has been already announced by Neumayr, this province was the autochthonous home of this family, and Neumayr’s opinion is strongly sustained by the remarkable series of species described from the Northeastern Alps by Geyer, Hauer, and others, and an especially fine series by Herbich from Siebenburgen. The Lytoceratidse are by no means absent from the faunas of the Lower Lias in Central Europe, though generally quoted as being found in the Middle and Upper Lias. Thus Amm. Drvani, Dumortier, and Amm. Salisburgense and Amm. altus of the same author, are apparently members of this family, found in the Oxynotus bed of the basin of the * Barrande, with all his knowledge and close study of the fossil Cephalopoda, has not been able to prove a single example. Those he has given are readily explained as morphological equivalents, and we haye found by the investigation of Bohemian specimens that the Nautili of the present time are entirely different from paleozoic forms. As soon as the nepionic and nealogic stages are studied and compared, they are found to be distinct. This is also true of his Gon. (our Celeceras) prematurum. * Geol. Verhiilt. d. Geg. nordw. vy. Achen-See. 112 GENESIS OF THE ARIETIDA. Rhone. The last two are closely comparable in aspect with species figured by Hauer from the Adneth limestones.1 In the same way, we should be disposed to regard the Mediterranean province as the autochthonous home of some genera of the Middle Lias, which appear here in association with the Arietide. The Arietide afford an excellent standard, since their genera and species have been found, with rare exceptions, only in the Lower Lias; and, so far as our knowledge now goes, the series of forms and cycles have a very complete and satisfactory aspect, indicating a history of progress and decline within the limits of that group of strata in the faunas of Central Europe. In the Mediterranean faunas, however, so far as known, only the rise of the group is recorded in the sediments and fossil remains, and its acme and decline are not clearly indicated. We have been accustomed to look upon the fauna of the Hierlatz beds as composed for the most part of degraded dwarfs, whose pecu- liarities or modifications were due to the unfavorable action of the surroundings upon migrants from other contemporaneous faunas of the Lower Lias. This seems to be the only theory which can account for the prevalent smaller size and more or less degraded aspect of many of the shells, when compared with their nearest allies in other locations. SUMMARY. The facts cited above, though far from complete, show that the series of the Radical and Plicatus Stocks, with the exception of the vermiceran series, were probably evolved in the Mediterranean province. The series of the Levis Stock had however a different history, since they probably arose in the basins of Cen- tral Europe. We therefore venture to differ in part from the eminent geologist and paleontologist Neumayr, who regards, if we properly understand his views, - the Northeastern Alps as the aldainic home of the whole of the Arietide. The sutures of all the Mediterranean forms of Psiloceras and Caloceras are, as figured by Wihner, more complicated, or, as we should say, more triassic than those commonly found in Central Europe; but we occasionally find a variety of Psil. planorbe, like that figured by Quenstedt? and by Wright,’ in which there is a close approximation to the outlines common in the Mediterranean province. After having written the above, we were extremely gratified to find precisely the same results with regard to the relation of caliphyllum and planorbe, but more fully and exactly stated by Neumayr, in his “ Unterster Lias” (p. 25). His conclu- sion, that planorbe is consequently a derivative of Psi/. caliphyllum, and is char- acteristic of Central Europe, while the latter species is equally characteristic of the Mediterranean province, is sustained by the fact that the sutures of calphyl- 1 The peculiarities of the senile whorls are similar to those of Oxynoticeras Lotharingum, and will lead to much confusion until the sutures and the young are fully known. [It is quite possible that our own con- clusion may be wrong in this respect, but the sutures of Salisburgensis and altus, Hauer, are Lytoceran, and the aspect of these compressed shells is very similar to that of those found in France, whose sutures are however unknown. The young are known only in Driani, which resembles some of the forms described by Herbich. 2 Amm. d. Schwab. Jura, pl. i. fig. 19. 3 Lias Amm., pl. xiv. fig. 1, SUMMARY. 115 dum become simpler with advancing age, and more like those of plunorbe, and by the scarcity of the latter, which, though found by Wahner,’ is declared to be rare. One of Wihner’s specimens was transitional to Hagenowi m its sutures, and this indicates that the province of the Northeastern Alps was the autochthonous home of caliphyllum, planorbe, and Hagenowi, and adds greatly to the probabilities in favor of Neumayr’s hypothesis. In Cad. Liasicum, Johnstoni, and nodotianum it is common to find varieties varying in the sutures between the Mediterranean and Central European extremes of modification, the latter being of course the most numerous in their own province and rare in the Northeastern Alps. The sutures: of Liasieum,? tortilis,? and nodotianus,s when contrasted with Quenstedt’s, Wright’s, and our own figures, give a good idea of the extent of variation, which is quite as great as in Psil. planorbe, if not greater. Undoubtedly these facts, and the nearer approximation in aspect and sutures of the Mediterranean forms of Psiloceras to Gymnites of the Trias, the genus we have always regarded as the probable ancestor of the former, are strongly in favor of Neumayr’s opinion that the forms of the Huropean province arose by — chorological migration from the apparently more ancient fauna of the Mediter- ranean province. The richer evolution of triassic forms in the Mediterranean province, as described and illustrated by Mojsisovics, can also be brought forward in fayor of this view. Nevertheless, it is not right to yield entirely to the fasci- nations of this opinion until there is positive proof that Psv/. planorbe or caliphyllum occurred earlier in this province than in Central Europe. With regard to the origin of Caloceras from this province, the facts are still stronger in favor of Neumayr’s view, but Vermiceras appears to have arisen in South Germany. With regard to the origin of Weehneroceras and Schlotheimia, it seems prob- able from the zodlogical evidence that they also arose in the Mediterranean provy- ince. The evidence is, however, geologically incomplete, since it is probable that Schlot. catenata occurred quite as early in South Germany. Weehneroceras, the series of connecting forms uniting Schlotheimia and Psiloceras in this same province, is not yet proved to be of as ancient origin as Schlotheimia itself, and this introduces an anachronism which requires additional facts for its explanation. Mosch ° has decided that the Lias to the west of the head-waters of the Rhine contains species peculiar to the Central European province. W. A. Ooster’s descriptions and figures of species confirm this conclusion, since he does not men- tion any novel species, though he describes twenty-one forms, representing more or less all the genera of the Arietidze.® Zittel” remarks that there is great resemblance between the Upper Lias in Provence and Lombardy. Mojsisovies,* in quoting these observations, says that 1 Verh. k. k. geol. Reichsans., 1886, p. 169. 2 D’Orb., Terr. Jurass., I. pl. xlviii. 3 Tbid., pl. xlix. 4 Tbid., pl. xlvii. 5 D. Jura Alpen d. Ost-Schweiz, 1872, p. 1. ® Cat. des Ceph. des Alpes Suisses, Denk. schweiz. Gesellsch. Naturwis., XVIII., 1861; see also Studer, Geol. d. Schweiz, II. p. 231, for similar views. 7 Central-Appenn., Geogn. pal. Beitr., Beneke, II. p. 174. 8 Dolomit Riffe Siid-Tyr. und Venet., p. 26. 15 114 GENESIS OF THE ARIETIDA. they raise the question whether the Mediterranean forms of the Swiss Alpine Jura may not have come by the way of southern France into the western Alpine region. j The very interesting and instructive essay of M. Dieulefait on the “Zone a Avicula contorta et Infra Lias dans le Sud-est de la France ”’ shows that in Proy- ence a southern and northern basin may be clearly separated. The southern or Mediterranean basin comprises a range of deposits reaching from the neighbor- hood of Toulon and Brignolles to Draguignan, Grasse, and Nice. The basin of the north and northwest, or of the Durance, encloses the valley of that river and the neighborhood of Castellane and Digne in the department of Basses Alpes. The basin of the Mediterranean possesses a series of beds identified as belonging to the zone of Avicula contorta, but there are no Ammonitine, and all the beds above these in the Lower Lias are absent. In the basin of the Durance, how- ever, a very complete series of lower lias beds, including a Planorbis and Angu- latus bed, has been described. M. Dieulefait has here traced the limits of the - Mediterranean province at a very important, and for our theory an essential locality. He has shown that the sharp division between the Mediterranean faunas and those of Central Europe, which, according to our conclusions, ought to exist along the boundaries between the basin of Italy and of the Rhone, can be actually traced in the field. Dumortier’s extensive observations in the valley of the Rhone and Collenot’s at Semur show the sudden spreading out by migration of forms of Psiloceras and Schlotheimia from South Germany into the Cédte d’Or at about the same time, and a somewhat later appearance of these radicals in the Rhone and North Ger- man basins, and possibly still later in England. It seems more likely also, from the two tables given above, that the species of Schlotheimia, Psiloceras, Caloceras, and perhaps Vermiceras, were migrants from the Cote d’Or basin to the Rhone, than that the reverse should have taken place. Coroniceras also thins out in this direction, whereas the genera haying their acme in the upper horizons of the Lower Lias, viz. Asteroceras, Agassiceras, and Oxynoticeras, are more abundantly represented, perhaps, than in the Cote d’Or. All the information obtainable with regard to the faunas of the Lower Lias in Switzerland indicates a general thinning out in numbers of species and varieties in that basin which, like the basin of the Rhone, lies to the south of the autochthonous zone. Emerson’s collection from Markoldendorf now at Amherst, Mass., and others we have seen, show that the fauna of North Germany was probably derived from South Germany, and this accords with Seebach’s conclusion, that a connection existed between the Hanoverian and South German faunas during the time of the deposition of the Lower Jura.2 There is considerable doubt whether the English species of Psiloceras and Caloceras came by the way of the Cote d’Or, or found this locality by independent migration. The former opinion is supported by the general fact that the English fauna does not contain an autochthonous series, nor does any radical species appear earlier in this basin than in those of the conti- nent; it is therefore probably a residual fauna, peopled by chorological migration. 1 Ann. des Sci. Géol., I. 1869, p, 473, pl. v. 2 Hannoverische Jura, p. 70. SUMMARY. 115 The prevalence of the geratologous forms of the different series in the highest beds of the Lower Lias indicates that this fauna, like those of the Swiss and Rhone basins, is also a residual fauna, but lying north instead of south of the zone of the autochthones. The only definite information with regard to the Lias faunas of the higher northern latitudes, which [ have been able to lay hands on, is the “I Sueriges Aldre Mesozoische Bildungen,” by B. Lundgren.’ Cor. Bucklandi and dzsulcatum are mentioned, and Cor. Sauzeanum,’ Agas. Scipionianum,? Agas. stri- aries,* and Arn. falearies are figured.? These indicate the presence of the Buck- landi beds in northwestern Sweden, but the fossils were in bad condition and not abundant in the number of species. Lundgren mentions, also, that these beds are underlaid by an unfossiliferous bed, which he thinks is probably the equivalent of the Planorbis and Angulatus beds of Central Europe. M. Hebert has, in his inter- esting paper, “ L’Age des Grés & Combustible d’Helsinborg et d’Hoganas,” * given proofs of the presence of the existence of the Planorbis and Angulatus beds in southern Sweden, but they contain no specimens of Ammonitine. It is well known that the Lias does not exist in Central Russia, and A. Pay- low, in his article on “ Russie, Esquisse Gologique,’’ gives an account of the deposits of the Jurassic, but mentions the Lias as occurring only in the Crimea and perhaps the Caucasus, and refers these to the fauna of the Mediterranean province, and not to Central Europe. Savi E. G. Meneghini, “Geologia della Toscana,” gives several lists of fossils from many distinct localities, among which are a number of the Arietide. Von Rath® quotes a list of fossils from Mene- ghini containing many Arietide, and he states that there are a number of new forms; but lists of names and descriptions of species are unfortunately not usually of value in such work as we are striving to do. Taramelli, in his mono- graph, “ Del Lias nelle Provincie Venete,”’*® describes and figures several species of Ammonitine. His Amm. Guibalianus is a true Oxyn. Gibali of considerable size, 300 mm. in diameter, and too involute for a specimen of Greenoughi. Arietites rotiformis is a young form of Cor. Gmuendense, or some such compressed shell ; it is assuredly not rotiformis if his figure is correct. Ar. obtusus is a true obdusum. Ar. stellaris is the adolescent form of Ast¢. stellare. All of these have the facies of the Northeastern Alps, except perhaps Gwali, which is new to us as occurring in the Mediterranean province. Sacco states, in his “ Lias della Valle Sturio di Cuneo,” ” that all the beds of the Lower Lias are present, and gives lists of fossils, including a supposed Psi. planorbe, several species of Schlotheimia, Ver. Conybeari, and a doubtful Cor. kridion, and Cor. Bucklundi and bisulcatum are said to be of good size and abundant near Pouriac. In “Lias Inferiore ad Arieti,” by C. de Stefani, it is distinctly stated, according to Geyer, that the Lower Lias of Italy is divisible into only two parts; ‘one which is similar to the Angulatus horizon, and yet contains the fauna of 1 Sueriges Geologiska Undersékning, ch. xlvii., Mollusk. 2 Pl. ii. fig. 5-7. B IPA. anti = Jel tis ie, OL © Jel, Tet, ie ES. 6 Ann. Sci. Géol., I., 1869. y 7 Annu. Géol. Universel, II., 1886, p. 302. S *«Geogn.-mineralo. Fragm. a. Italien,’’ Zeitsch. deutsch. geol. Gesellsch., XX., 1868, p. 320. ® Atti dell’ Instituto Veneto, ser. 5, V., 1880, Appendix. 10 Boll. del R. Comitato Geol. d’ Italia, XVII., 1886, p. 15. 116 GENESIS OF THE ARIETIDA. Spezia, and another higher horizon, which is supposed to be the equivalent of the Bucklandi horizon. This last is said to contain Cephalopods representing all the later faunas of the Lower Lias, and’some species are quoted as being referable to the Middle Lias. Canavari, in his “‘ Fauna der unteren Lias von Spezia,” so frequently quoted above, states that the fossils occur in a single zone, which does not admit of sub- division, though it was carefully investigated, layer by layer, by Cocchi. He states also, that it is unquestionably the lowest of the lower lias sediments in Italy, and comprises all the horizons except those of Planorbis and Oxynotus. He considers that the fossils have.closer affinities with those of the Mediterranean province than with those of Central Europe, a fact which seems to be established. The species of the Lytoceratidze and of Amaltheus, ete, which are supposed to be anachronic and to indicate a fauna derived from the Middle and Upjer Lias, appear to us to be found in their appropriate positions, like those of the Northeastern’ Alps. They may be either the radicals of the similar forms which occur in the Middle and Upper Lias of the Central European faunas, or morpho- logical equivalents, or pathological specimens.' This is also Canavari’s opinion with relation to some forms, since he expressly states that the segoceran species, as he calls them, are the immediate forerunners of Microderoceras. The fauna of the Rhone basin is almost exclusively composed of species having a Central European aspect. There are, it is true, some slight mdications, in the presence of three species of Lytoceratidz in this basin, that the migrants may have come this way on their march into-Central Europe, but there are no supporting facts with which we are acquainted. The absence of the Planorbis horizon, or at any rate its sporadic appearance in Italy, and the absence of Ammonitinz in this horizon of southern Provence, are very serious difficulties in the path of a sup- posed southern track of migration. The evidence, so far as known, seems therefore strongly in favor of the view, that during the time of Planorbis and Caloceras, and perhaps earlier in the Angulatus horizons, the stream of migration flowed south and westerly from the Northeastern Alps into Italy, while another from the same basin directed itself westerly along the then existing coast lines into the basins of South Germany and the Cote d'Or, and the species were distributed thence into the basins to the north and south of these two, in the province of Central Europe. In South Germany and the Cote d’Or the conditions became favorable during the time of the Angulatus horizon for the evolution of Vermiceras among the descendants of the Plicatus Stock, and for the origin of Coroniceras, Arnioceras, and Agassiceras of the Levis Stock. Asteroceras arose later in these same faunas in the Upper Bucklandi beds, and Oxynoticeras probably even still later, though here the series is evidently older than the date of its first appearance. The migrations of these genera spread the forms to the east into the faunas of the Northeastern 1 The figures of Amaltheus given by Canavari in his last work, ‘‘ Fauna del Lias inf. della Spezia, R. Comit. Geol. d’ Italia,” III., Pt. IT. pl. vi., are certainly startlingly similar to Amaltheus, but such resemblances in forms of widely distinct series are not uncommon. See the pathological case figured on Plate X. Fig. 19 of this memoir, and others quoted in the descriptions of the species. SUMMARY. 117 Alps, and thence they passed southerly into the Italian basin. Migrants also passed in all other directions into the residual basins to the north and south of the basins of the Céte d’Or and South Germany, in the province of Central Europe. While these faunas in the Northeastern Alps and Italy became analdainic faunas so far as the Arietidze were concerned, they were aldainic faunas for some other groups, like the Lytoceratidz, and also very likely for the Liparoceratide, Deroceratidse, and possibly other families. These mixed faunas, which have been deemed such sources of confusion, are in reality the most instructive, and will enable us to trace both chronological and chorological migrations with greater security, if the views here advanced are correct. Table V. shows that there are but two examples of what Neumayr calls cryp- togenous types in Central Nurope, species appearing suddenly without apparent ancestors, Schlot. catenata and Psil. planorbe, var. leve. Schlot. catenata, liowever, cannot be called an unquestionable eryptogenous form in the Northeastern Alps. It is in that basin connected by intermediate forms, as stated above, with Psilo- ceras, and it is therefore probable that in course of time the geological evidences which are now confusing will be brought into accord with the paleozodlogy. Psil. planorbe is a radical derived from Psi. calphyllum, or else from pre-existing triassic ancestors, and the absence of a complete series connecting it or Psi. caliphyllum with Gymnites of the Trias is evidently due to the absence of an equally complete series of formations. That the intermediate species might have been deep-sea forms, and therefore not represented in the rocky strata now exposed, as supposed by Neumayr, is an admissible explanation, Newberry’s hypothesis * of the retirement of the sea is, however, equally supposable, and has the additional recommendation of explaining the absence both of intermediate forms and of the sediments. Newberry thinks that the presence of intermediate Inks in paleozodlogice history, and their absence from localities so far explored, are explicable on the supposition that the chain of the rocky deposits is incom- plete in those localities, and that the sea had retired from them carrying with it the threads of life. The missing links of the record were then evolved in other places, but not brought back by the return of the ocean to its former shores. This seems to us more in accord with what is already known of the merely frag- - mentary aspect of the geologic record in any one region, the occasional discoy- ery of the absent leaves of the record in other places, and the want of absolute synchronism between the strata of Europe and those of America. That Psi. planorbe was a littoral form, as well as its congeners, can hardly be doubtful, since, besides the facts quoted above, they are found associated in the same series of layers with bones of saurians and even remains of insects. in Eng- land. The remarks of Martin and other authors, quoted above, upon the charac- teristics of the lumachelle in the Céte d’Or, and the broken aspect of the shells of Ammonoids compared with those of swimmers like the Nautiloids, as stated by Terquem, in the department of Moselle, the opinions of Tate and Blake, and ? Circles of Deposition in American Sedementary Rocks, Proc. Am. Ass. Ady. Sci., XXII., 1873, pp. 185, 189. 118 GENESIS OF THE ARIETIDA. the great abundance in which Ammonoids occur as contrasted with Nautiloids, are all in favor of the conclusion that they were structurally rostrated, creeping animals, which necessarily followed the shore lines in their migrations. Fraas takes the view that the Suabian Lower Lias was as a whole, when compared with the synchronous strata to the west and north, a deep-sea formation, and cites the absence of sandstones and coarse deposits, the small Lamellibranchs, and Brachiopoda. It is very evident, however, that whatever the bathymetrical differences of the South German basin, and however far removed from the ancient shores now represented by the Black Forest and the Vosges, the sur- roundings were not sufficiently distinct to make any marked differences in the Arietidze of this basin. We have noted above the occurrence in Peru of Cal. Orton’, a form having close resemblance to a species of the Northeastern Alps, and the apparent iden- tity of other species with those of Central Europe, the forms found at Vancouver's Island and in California, etc., show that on this continent the faunas possessed a mixed character. The paucity of the development both geologically and paleon- tologically of the Lower Lias is in accord with the similar deficiency of this stage in the analdainic basins of northern Europe, India, and Italy. There is another fact in this connection, which strikes us as very remarkable, — the absence of any absolutely new types of Ammonitine. So far as explorations have gone, not a single species indicates the evolution of any widely distinct family or genus from those found in Europe. Thus, although not able to produce any satisfactory evi- dence that all the faunas throughout the world during the lower lias age were more or less analdainic faunas derived from the zone of the autochthones of the Arietidze in Europe, the evidence is sufficient to make such an opinion worthy of the attention of students of geology and paleontology. The view expressed by Neumayr, that the Cephalopoda are exceptional in respect to the rapidity with which their modifications probably took place, seems to us erroneous. There is no greater aspect of pliability in this than in other types, when accurately classified. When, however, we assemble within the same family species of the Lytoceratinee and Ammonitine, or in the same genus forms of entirely distinct stocks without sufficient reference to their genetic history, then of course a belief in the polygenesis of the progressive series, and in an exceptional tendency to modification, becomes essential in order to explain the heterogeneous aspect of the groups. We think, however, that even the most variable families of Cephalopoda are not, as a rule, any more variable than the Unionide, Ostreade, or Hippuritidee, among Lamellibranchs, or the Planorbide, Vermetidze, etc., among Gasteropoda, and many other groups that might be mentioned. The expansion of the whole series of forms of Psiloceras, Schlotheimia, and Weehneroceras in the Northeastern Alps, and the apparent rapidity of chorologi- cal migrations and changes and introduction of new series, the equally sudden 1 We desire to call attention here to the fact that we have admitted the polygenetic derivation of retro- gressive types like Baculites, ete.; but this in no manner commits us to the doctrine of polygenesis for any of the progressive types. So far as we know, these are monogenetic in mode of origin, Bi SUMMARY. 119 _ expansion of the Arietide in the Bucklandi zone of Central Europe, the rapidity with which the forms of the still later beds must have come into being in order to be presented in a body, as in the Tuberculatus beds of the Cote d’Or, and the limited thickness of the beds, are all against the supposition that it required vast periods of time for a species to become modified and give rise to series of distinct forms. Hither the species of the Arietide had time enough during the deposi- tion of the Planorbis, Angulatus, and Bucklandi beds of the Lower Lias to spread themselves over the entire area of modern Europe, and generate from one form all the series described above, or else the same species and genera had invariably distinct centres of origin in the different basins. One might support the latter view and favor polygenesis even in this extreme sense with considerable show of reason, if there were not such a mass of evidence in favor of migration, some of which we have given above. If there were space, we could add examples from the researches of various welt known zodlogists upon the migrations and modi- fication of species in modern times, both along the coasts and over the land. The more striking examples are, however, quite well known, and hardly need to be dwelt upon. 120 GENESIS OF THE ARIETIDA. V. DESCRIPTIONS OF GENERA AND SPECIES OF ARIETIDZ. RADICAL STOCK. FIRST, OR PSILOCERAN BRANCH. PSILOCERAS. HELL smooth, plicated or with fold-like pile in some subseries. The abdo- men is rounded, or with smooth median zone, never channelled or keeled. Whorl in section is compressed, helmet-shaped. ‘The sutures are similar in pro- portions and outlines to those of Caloceras. This is shown in the broad abdomi- nal lobe and large siphonal saddle, the equality in length and size of the abdomi- nal and lateral lobes and saddles, their leaf-shaped marginal digitations, and the number and inclination posteriorly of the auxiliary lobes and saddles. The living chamber is one, or more than one, volution in length, and is shorter in the young than in the adult stages.’ Senility is indicated by increasing con- vergence of the sides, and the loss of plications,’ but a subacute abdomen, such as appears in the old whorl of Weehneroceras, is never present. The completeness and accuracy of Wihner’s illustrations and descriptions, which enable one to study all the stages of growth in some species, has tempted us to suggest the existence of three subseries in this genus. (1.) The first contains smooth shells, typical helmet-shaped whorl, and an old age in which a subacute whorl is not yet re- corded in any species. (2.) The second contains plicated shells exactly similar in form, but the folds numerous and regular, and in some species figured by Wihner these cross the abdomen with a forward bend. They are, however, not true pile, and, so far as we know, they do not become depressed along the median zone as in Wehneroceras. (3.) The third contains shells having psiloceran forms but flattened sides, and often plicated as in the second subseries, though the Psil. Hagenowt is smooth. We regard this subseries as of doubtful utility, but do not know how to dispose at present of the forms it contains. 1 Throughout this chapter there is no attempt to give a complete synonymy of any one species. The references given under each name are only those which were considered essential to settle the applica- tion of the specific name and the range of the forms to which it was applied in this memoir. The localities given are those of specimens in the collections of the Museum of Comparative Zodlogy. 2 Quenstedt, Amm. Schwab. Jura, pl. i. fig. 6, shows a nealogic stage in which this chamber is not quite half a volution in length. Wihner takes note of this, (Unter. Lias d. norddst. Alpen, Mojsis. et Neum., Beitr., 1V., 1886, p. 185,) and states that in one example of Psil. planorbe from Wiirtemburg observed by him the living chamber was only two thirds of a volution in length, and suggests the same opinion with regard to the shorter living chambers of the young. 3 Quenstedt figures what may be a fragment of an old specimen of Psil. planorbe, Amm. Schwab. Jura, pl. iii. fig. 1, and Wiihner has figured several old specimens in Unter. Lias, Mojsis. et Neum., Beitr. FIRST, OR PSILOCERAN BRANCH. 121 First AND SECOND SUBSERIES. Psiloceras planorbe, Hyarr. Var. leve. Plate I. Fig. 1-4. Summ. Pl. XI. Fig. 1; Pl. XII. Fig. 1. Amm. planorbis, Sow., Min. Conch., V. p. 69, pl. ceeexlviii. Aigoc. planorbis, Wrigut, Lias Amm., p. 308, pl. xiv. fig. 1-4. Amm. psilonotus levis, QUENST., Die Ceph., p. 73, pl. i. fig. 13; Amm. Schwab. Jura, pl. 1. fig. 1-7. Amm. Sampsoni, Porti., Rep. Geol. Londonderry, ete., p. 138, pl. xxix. a, fig. 13. Psil. planorbe, Hyatt, Bull. Mus. Comp. Zool., I. No. 5, p. 73. Localities. — Whitby, Watchet, Montloy, Semur, Rudern, Nellingen, Balingen, Neuffen.1 This remarkable form is a somewhat flattened discoidal and perfectly smooth shell in its typical adult form. The young are often plicated. Var. plicatum. Plate I. Fig. 5, 6. Summ. Pl. XI. Fig. 2. Amm. psilonotus plicatus, QuENST., Amm. Schwab. Jura, pl. 1. fig. 1-14 (mot fig. 8, 13). This shell differs from variety /eve merely in having immature pile or folds in the neologic and ephebolic stages. There is therefore the most gradual and hardly perceptible gradation from the preceding variety to this form. The septa of both are exceedingly variable. The marginal digitations may be either very shallow, as in the Arietidee generally, or they may be foliaceous and complicated, as in the radical series. The lobes and saddles may also vary exceedingly in size and proportions; some species have deep and narrow saddles with long broad lobes, as in the radical series, while others, more like the typical Arietide, have shallower, broader saddles, and shorter, more pointed lobes. In the collection at Semur there are forms from Saulieu identical with the South German, which when compared with raricostatum and Johnston, show closer approximations than any specimens seen elsewhere. The Bristol collection contains undistorted specimens of this species from Cotham, and in Dr. Wright’s collection from Whitby the piicatus variety is labelled Amm. erugatus, Bean. The connection with the flattened Watchet specimens of planorbis, Sow., can be clearly made out by the large tablet in the British Museum, containing about one hundred and fifty specimens. Of these, perhaps ninety exhibit folds like those of plicutus and erugatus. The largest on this slab is from 60 to 80.5 mm. in diameter. These large specimens are not equivalent to Cad. Johnston, as Oppel supposed, but to plicatus. Hrugatus seems to be a dwarfed form with the folds often developed very strongly in the young,’ and the shell has fine strie of growth, as in Agas. striaries, Plate IX. Fig. 14, 15. In the Museum of Comparative Zodlogy the series is complete from /eve to var. plicatum, as figured by Quenstedt in “ Der Jura,” and in another direction to the var. of planorbe from Semur.? This is a slightly plicated form, having the sides of the whorls broader 1 These localities also include var. plicatum. 3 See Pl. i. fig. 5, 6. 2 Amm. erugatus Bean has only the young plicated, resembling in this respect var. /eve. It is however always a small form or dwarf. 16 122 GENESIS OF THE ARIETID/. than usual, and the involution slightly increased, a modification which is also sometimes present, though less marked, in erugatus. Wiihner found what he claims’to be Ps?. planorbe at Pfonsjoch in the Pla- norbis bed. These were small specimens, measuring 15-40 mm. in diameter, and one of them is said to be similar to Hagenowi.' In the same work he figures the following discoidal shells of the smooth subseries: Psz/. polycyclum and cali- phyllum, Plate XV., and Psil. pleurolissum. Neumayr, in the Unterster Lias,’ gives Psil. planorboides, a more involute, smooth species of this series. Planorboides appears to lead into two much more inyolute and compressed species figured by Wiihner in the same work, Psd. Atanatense and mesogenos* Both of these are devoid of true pilee, and possess only senile fold-like pilations.’ SECOND SUBSERIES. Psiloceras longipontinum, WAuyer. Amm. longipontinus, Orr., Pal. Mittheil., p. 129, pl. xh. Psil. longipontinus, WAuNER, Unt. Lias, Mojsis. et. Neum., Beitr., IV. p. 196. Egoc. Clausi, Neum., Unterst, Lias, Abh. k. k. geol. Reichsans., VII, pl. iii. The original of this species in the Museum of Stuttgardt has considerable like- ness to Ps?/. planorbe, var. plicatum. Oppel seems to have considered it one of the schlotheimian series.’ The open umbilicus, straight folds in place of true pilee, keelless abdomen, and helmet-shaped form of whorl, show it to be a member of the psiloceran series. The sutures,’ as figured by Oppel, exhibit the strong psilo- ceran affinities of the species. In his specimen the last whorl has become smooth on one side, and the pilz nearly obsolete on the other, thus mdicating the approach of senility, though the shell is but 95 mm. in diameter. The pile begin to obsolesce posterior to the last septum. The living chamber is nearly one volu- tion in length, though still incomplete. An empty cast in the Semur Museum from Ruffy undoubtedly belongs to this species; it is 155 mm. in diameter, and the last whorl is smooth, showing its great age. There are specimens in the collec- tion at Munich labelled Amm. Roberti, Hauer, locality Filder, and Amm. Oeduensis, 1 Unter. Lias, Mojsis. et Neum., Beitr., IV. p. 136. 2 Tbid., Il. pl. xxvii 3 Abhandl. geol. Reichsans., VII. pl. iv. 4 Op. cit., III. pl. xxvi. 5 We have figured only the most involute of this smooth series on Summ. PI. xi. fig. 15. 6 The closeness of the parallellism between some of the forms of Psiloceras and some species of the Lytoceratidz is such as will be likely to cause considerable confusion unless great care is taken in studying the species. Comparison of such forms as Amm. Petersi, Hauer, Ceph. nordost. Alpen, pl. xxi. fig. 1-8, Lyt. Petersi, Herb., Széklerland, pl. xx., Lyt.? Driani, sp. Dumort., Etudes Pal. du Basin du Rhone, and Lytoc. (Amm.) Roberti, Hauer, Capric. oesterr. Alpen, pl. iii., will show that without close study of the sutures and young no separation can be made with certainty. In fact, in identifying Driant in the absence of figures of the sutures as a form of Lytoceras, we have been led by the geological position and size, which accord better with Lytoceras than with a species of Psiloceras. It is possible that in doing this we are illustrating these remarks in a forcible manner. See also in this connection the forms of Rhacophyllites and Phylloceras figured by Canayari in his ‘¢ Fauna des unteren Lias von Spezia,’’ pl. ii. 7 The sutures figured by Portlock, as well as the form of the section (b) of his Amm. Sampsoni (fig. 13 ¢, not fig. 13 a), suggest longipontinus, and may indicate the presence of this form. or transitional varieties, in the English basin. FIRST, OR PSILOCERAN BRANCH. 128 Despl. d. Champ., locality Blumenstein am Thuner See. One of the specimens from Filder shows the exact aspect and markings of Psi, planorbe, but has the form of longipontinum. Though it is somewhat difficult to judge from a figure, nevertheless, 44y. Clausi, Neumayr, very closely resembles Psi. longipontinum, and we have considered it to be a variety of this species with somewhat stouter whorls than the normal form. It is also a large aged specimen, and according to Neumayr came from Wiirtemburg. Quenstedt referred this species, in his “ Ammoniten des Schwiibischen Jura,” to Cal. laquewm. His comparisons were evidently made with the old whorl of laqueum, and, as this has no keel, and is smooth or with obsolescent pile, it is of course very like the adult stages of Psél. longipontinum. Nevertheless, both the young and adult stages of daquewm are easily distinguished from the same stages of longipontinum. Quenstedt’s figure shows the length of the living chamber to have exceeded one volution. Tate and Blake’s citation of this species from the Angulatus bed? is likely to mislead. Their species is, as figured, a diseased Caloceras, or poorly drawn species of Schlotheimia with pile crossing the abdomen, but certainly not, as named by them, dongipontinus. Species of the second subseries figured by Neumayr in the work quoted above are as follows. Psi/. eryptogonium, Plate VL, is discoidal. Ps¢/. majus and Gernense, Plate V., are slightly more involute shells. Wihner, in Volume IV. of the work above quoted, figures Psi. sublaqueum, Plates XV., XVI., Psi. crebri- cenctum, Plates XVI., XVUL, Psil. pachydiseus and polyphyllum, Plate XVII., all discoidal shells. This subseries and the preceding agree closely with the western European forms except in the involute species. Wiihner also figures, in Vol- ume III. of the same work, Psi/. Berchta and aphanoptychum, Plate XXIII, which are discoidal, and Psi. pleuronotum, Plate XXV., caleimontanum, Plate XXIV., and Kammerkarense, Plates XXIV., XXV., which are more involute and compressed. THIRD SUBSERIES. Psiloceras Hagenowi, Wiuver. Amm. Hagenowi, Dunx., Paleontogr., I. pl. xiii. fig. 22, pl. xvii. fig, 2. Amm. Hagenowi. Tera. et Prnv., Lias Inf. de Est de la France, Mém. Soe. Géol., VIII. pl. i. fig. 3, 4. Amm. Hagenowi, Quenst., Amm. Schwab. Jura, pl. i. fig. 18. Psil. Hagenowi, Wkuner, Unt. Lias, Mojsis. et Neum., Beitr., IV. p- 196. The form of this shell approximates to that of Psil. planorbe, var. leve, but the sutures are more widely distinct, and degenerate in outline. In Terquem and Piette’s figure they resemble quite closely the sutures of Popanoceras Kingtanun and antiquum, Goniatitinee of the Dyas. The lobes of that figured by Quenstedt are not so coarsely dentate, and approximate more closely to the sutures of Psil, 1 Yorkshire Lias, p. 273, pl. v. fig. 4. ? On Summary Pl. xi., outline figures have been given of the principal forms, aphanoptychum, fig. 11, and Kammerkarense, fig. 12. 124 GENESIS OF THE ARIETIDA. plunorbe. The saddles sometimes have entire margins, as in some Ceratitinee of the Trias. Neumayr’s Hagenowi, in his “Unterster Lias Nérdalpen,”?! is not a true Huagenowt, if the sutures are correctly drawn. Such facts and the remark of Wiihner quoted above (page 113) show that Hagenowi is probably a dwarfed deformation of Psi. planorbe, which is likely to occur in any locality, and has an’ independent existence as a race or species only in certain basins where it is abundant. It seems to indicate, wherever it appears, that Psi. planorbe has there been subject to unfavorable conditions. Neumayr in the work above quoted, Plate IV. Fig. 1, gives a Psil. (Ayoc.) Naumann, a good-sized species with numerous folds, compressed slightly conver- gent sides, and a rounded smooth abdomen, exactly the form and characters of his Hagenowi, except that it is more decidedly plicated. The smaller Psi. (AZyoc.) crebrispirale, Ibid., Plate V. Fig. 4, is probably the young of this shell. The sutures have complicated margins, as in other shells of this province, and are not similar to those of Hagenowi. We place it here until its exact affinities can be settled by the study of a series, or of the young. Migoceras Struckmanni,? Ibid., Plate VI. Fig. 5, as remarked by this distinguished authority, is a unique survival of triassic forms. It resembles the flat-sided whorls in Tirolites, and even certain earlier forms, like Popanoceras. It may be provisionally associated with this series until the sutures are known, since the shell is smooth, and similar to that of Ps¢/. planorbe. This series should be care- fully studied with ample materials. It may be that confusion exists between some forms now supposed to be true Psi. Hagenowi and some triassic forms still surviving in the Lias. Canavari, in his “ Fauna del Lias Inferiore di Spe- zia,’ 1888, Plate VII., has figured a series of what appear to be true Tropites. These are very close congeneric forms of this triassic genus, and in our opinion should be referred to Tropites itself? This gives greater force to the suggestion made above. Canavari, in his “ Fauna des unteren Lias von Spezia,” gives some interesting forms of this genus. Psi/. (goc.), Plate XIX. Fig. 2, 4, 5, is a plicated form similar to pleuronotum. Psil. pleuronotum, Plate XIX. Fig. 3, may possibly be the same as Psil. calcimontanum, as stated by Wihner, but it is a dwarf, like most of the species from this locality. Psi. (Mgoc.) Portisi, Plate XIX. Fig. 6, appears to bear a similar relation to Ps?l. mesogenos. Wihner, however, considers it iden- tical with the young of his Psd. Kammerkarenses 1 Abhandl. geol. Reichsans. Wien, VII. pl. ii. 2 Wahner’s Psil. Struckmanni, Unt. Lias, Mojsis. et Neum., Beitr., IV. p. 196. 8 The survival of characteristic triassic forms in the Jura shows that the connections between these two systems are closer than has been supposed, and gives support to opinions advocated in the chapter on Geological and Faunal Relations, and adds another group to the three already noted, Psiloceras, Lytoceras, and Phylloceras. These facts demonstrate that no insuperable barrier arrested the migration of forms and the continuity of the faunas in time. See remarks on Tropites in note to page 154. 4 Paleontogr., XXIX., and also Mem. del. Car. Geol. d’ Italia, III., 1888. SECOND, OR SCHLOTHEIMIAN BRANCH. 125 TMA\GOCERAS. Tmaegoceras latesulcatum, Hyarr. Amm. latesulcatus, Havrr, Ceph. d. Lias. d. Nordostl. Alpen, pl. ix. fig. 1-3. This extraordinary form, found in the red limestones of Adneth, has a combi- nation of characteristics altogether distinct from that of any other species. The form of the whorl, its smooth shell, and the discoidal mode of growth, are purely psiloceran. The sutures are, however, arietian, and more like those of Caloceras than typical Psiloceras or those of any other genus. We are not aware of its having been found elsewhere than in the Mediterranean province. Hauer appears to think that its affinities may lie with the Arietide, and that is also our opinion, but until the young have been studied it cannot be classified. Tmaegoceras levis, Hyarr. Ariet. Levis, GeyeR, Ceph. y. Hierlatz b. Hallstadt, pl. iii. fig. 10. This is a smooth, keeled, and channelled discoidal form like the preceding, but dwarfish, like other species of this locality. PLICATUS STOCK. SECOND, OR SCHLOTHEIMIAN BRANCH. The living chamber is of uncertain length, though Quenstedt gives it in his « Ammoniten des Schwibischen Jura” as possibly a volution in length in Schlo- theimia. The shell is involute in some forms. The whorl is flattened laterally, and in old age became subacute. A smooth median zone or channel was formed on the abdomen by the suppression of the pila, which were continuous across the abdomen in the preceding nealogic or ephebolic stages. There are no genicule, though the pilz are very completely developed. The forward bend is necessarily gradual, the whorl never having a sufficiently quadragonal form for the forma- tion of abrupt bends or genicule on the edges of the abdomen. ‘The sutures resemble those of Psiloceras and Caloceras. W ZH HNEROCERAS.? The adult has a smooth median zone along the abdomen. The pile, so far as the young are known, cross the abdomen during the earlier nealogic stages, and this character is retamed throughout the adult stages in some species. The smooth zone is really an incipient channel, formed subsequently by the resorption of the pile. This process may take place either in the later nealogic, ephebolic, or senile stage, according to the species. In old age the pile tend to degenerate into folds, and 1 Tunyos, a furrow. 2 Dedicated to Dr. Frantz Wahner, as a token of respect for his remarkably accurate and instructive researches upon the Arietide. 126 GENESIS OF THE ARIETIDA. become wider apart, the abdomen narrower, and the whorl consequently much compressed and subacute. No proper quadragonal whorl is formed during the growth, and therefore the senile outline of a section of the whorl is not trigonal, as in the senile stages of shells of other branches having a flattened abdomen and a keel in the ephebolic stages. All the species have true pilx, though these are not prominent, and the earlier nealogic stages resemble adult specimens of the second subseries of Psiloceras, in which the folds are well developed and cross the abdomen. We cannot distinguish either this genus or Schlotheimia, or Caloceras, from Psiloceras by means of the sutures. Psil. sublaqueum, Wiih., and other species of Psiloceras having plications which cross the abdomen until a late stage of growth, are not distinguishable until they are nearly full grown from some discoidal forms of this series. Waehneroceras subangulare, Hyarr. Amm. subangulare, OprEL, Paleontolog. Mittheill., p. 130. We have referred the species to this genus entirely upon the information derived from notes made before Wehneroceras was separated. It will be seen, however, that no species of Schlotheimia has young which remain similar to Psiloceras for such a prolonged stage as in Weehneroceras. One of the types of Amm. subangularis, Oppel, from Kalthenthal, in the collec- tion at Munich, has a form similar to that of Psid. planorbe, and pilations which cross the abdomen. The young is also a pure planorbe until over 14 mm. in diameter. Another specimen from Filder, which we have referred also to this species, has curved and close-set pile, and the form and smooth abdomen of planorbe (not channelled at all) until over 26 mm. in diameter; then the pilx begin to cross the abdomen. This last specimen was named Amm. planorbis by Oppel. There are also specimens from Hammerkhar, formerly referred by us to subangulare, which may be distinct. They certainly possess characters which were noted by us as intermediate between this form and true angulata, and one of them has a very peculiar old whorl, and may be a caloceran form.) Waehneroceras tenerum, Hyarr. 4Egoc. tenerum, NeEuM., Unterst. Lias, Abh. k. k. geol. Reichsans., VII. pl. iii. fig. 4, 5. Psil. tenerum, WAn., Unt. Lias, Mojsis. et Neum., Beitr., IV. p. 198. This form, described by Neumayr as occurring in the Northeastern Alps and also in Central Europe, at first seemed to us identical with Weh. subangulare. Neumayr remarks that, though the young are so similar, the adults are separable, and we have upon his authority held it to be distinct. He also looks upon this species as very closely allied to Psiloceras, and to be a transition form from the latter to Schlot. angulata. Species of this series have been figured by Wihner in Volume III. of his “Unteren Lias” as follows: Woh. Paltar, Plate XXI., and Rahana, Plate XXIIL.; 1 Proc. Bost. Soc. Nat. Hist., XVII., 1874, p. 18. SECOND, OR SCHLOTHEIMIAN BRANCH. 127 and in Volume II. he figures Wek. extracostatum, Plates XIV., XVI., and Panzner‘, Plates XV., XXI. The figure of extracostatum shows an old whorl which is acute, but not involute. Among discoidal shells, Wah. circacostatum, Plates XV., XVI., curviornatum, Plate XVI., and haploptychum, Plate XVII., show that the whorls of their earlier nealogic and adult stages are without channels. Wéeh. anisophyllum, Plate XIX. Fig. 1 a, shows a very old stage with subacute trigonal whorl, and pile replaced by folds. Woh. megastoma, Plate XVIII. Fig. 2, 3, shows ear- lier nealogic stages with pile continuous across the abdomen in the adult and senile stages. Woh. euptychum, Plates XVUIL, XX., stenoptychum, Plate XX., latimontanum, Plate XX., and diploptychum, Plate XXI., also belong to this ge- nus. The last two are senile specimens, with subacute outer whorls, and all the above are discoidal shells exhibiting transitions from Psiloceras to Schlo- theimia. ‘ There are, however, involute forms in this series also figured by Wiihner in the same work, but in Volume IV. These are Wah. Gudoni, Plate XXVI. Fig. 3 a, b (not Fig. 7), and Weh. Emmrichi, Plate XXVI"' We doubt whether either of these involute forms can \ be regarded as transitional to Schlotheimia, as sup- posed by Wihner. The results of our work upon the nealogic stages and their meaning in Schiot. catenata, and all mA ai re other species, show that series arose only from dis- The, W-22,— Wiema Prom fin Bear, coidal shells, and probably never originated from EN Cie note ee AO WE : Emmrichi, after Wihner, showing the the compressed and involute forms. These are involution of this species. The charac- k : : . : " teristic pile and channelless abdomen of themselves invariably discoidal and less compressed 445, genus are also noticeable in these in their own young, showing them in every case figures.’ to have been derived from shells having depressed abdomens and discoidal whorls. D « Canavari, in his “ Unteren Lias von Spezia,” * describes and figures dwarfs or the young of Woh. (Agoc.) Emmrichi under the name of Gurdon’. Wahner thinks that Canavari’s forms (Plate XVIII. Fig. 14, 15) are referable to his Gadoni, and Fig. 16 to be identical with his Emmrichi. The last is to us a very remarkable form, since it possesses continuous lateral and abdominal constrictions. SCHLOTHEIMIA. The form varies in this genus from discoidal to involute, but the umbilici are never entirely covered in. The whorls are usually flattened more or less on the sides, and the abdomen depressed. In the nealogic stages this form is common, 1 We have given outline figures of Wah. curviornatum, Summ. PI. xi. fir. 7, haploptychum, fig. 8, toxo- phorum, fig. 9, and Emmrichi, fig. 10. 2 This figure, according to Wahner, is poorly drawn, the last volution too narrow, the umbilicus too open. It, however, exhibits the general aspect of involute forms in this series, and we have retained it with that purpose in view. : 3 Paleontogr., XXIX., and Mem. del. Carta Geol. d’ Italia, III., 1888. 128 GENESIS OF THE ARIETID. but in the adults of involute species the whorl is necessarily more compressed. The compressed stage occurs very early in the most involute species, the flatten- ing of the sides and the depressed abdomen being omitted. ) © Neumayr, 5, 6, 21, 29, 32 nole, 33, 42. 56, 60, 61, 85, 86, 88, 89, 90, 108, 111, 112, 113, 117, 118, 122, 123, 126, 129, 184, 137, 189, 140, 147, 150 151, Neyada, 172, 173. Newberry, 117, 151. Newbold Quarries, 155. Nice, 114. Norites, 3, 4. North America, 86, 87. Northeastern Alps, xi, 55-57, 60, 61, 81, 85, 89, 90, 94-102, 106, 108, 110-113, 115-118, 126, 136, 137, 139, 140, 151-153, 159, 202. North English Basin, 88. North Europe, 118. North German Basin, 88, 92, 93, 102, 108, 139. North Germany, 92, 93, 94, 96, 97, 98, 99, 100, 101, 102, 108, 114, 129. North Peru, 157, 163, 169. Northwest Germany, 92. Nostologic Stages, 20, 28, 32, 37, 46, 51, 183. Nurtingen, 187, 182. Obtusus Bed, 107, 109, 188, 196, 202, 206, 207, 208, 210. Oelschiefer, 163. Oestringen, 141. Ofterdingen, 209. Olcostephanus, 23, 33. Olecostephanus Gravesianus, 23. Ontogeny, x. Ooster, 90, 113. Ophidioceras, 36, 47. Ophioceras Johnstoni, 144. kridioides, 171. raricostatum, 145. Oppel, 85, 97, 109, 110, 121, 122, 126, 138, 144, 149, 150, 159, 160, 167, 175, 188, 196, 209, 218, 218. Oppelia hecticus, 24. steraspis, 51. Organic Equivalence, viii. Origin of Differentials, 48. Floral Structures, 50 note. Variations, according to Weissman, 43. Orthoceras, 1, 15, 26, 54, 37, 39, 46. Orthoceras crotalum, Hall, 35 nole. docens, Bar., 34. fusiforme, Hall, 35 note. politum, 11. truncatum, 11. unguis, 11. Orthoceratide, 12, 13, 20, 25, 35, 36, 48, 49. Orton, 163, 169. Osterhornes Mts., 90, 93, 95, 96, 97, 100, 109. Ostreadze, 118. Ovum (monoplast), 8. Owen, Richard, 8. Oxynoticeran Branch, 214, 215. Series, 40, 69, 104, 106, 107. 0 INDEX. Oxynoticeras, 50, 55, 54, 47, 48, 54, 55, 71, 77, 78, | Placenticeras placenta, Meek, 31. 80, 82, 83, 84, 101, 106, 107, 108, 109, 114, 116, } Planicosta Bed, 105, 209, 213. 148, 214. j Oxynoticeras Aballoense, 220. Bavigneri, 220. Castagnolai, 216. Collenoti, Geyer, 213. Greenoughi, 69, 213, 214, 216, 218. Guibali, 69, 83, 115, 214, 215, 218, 219. Guibalianum, 218. Janus, 221. Lotharingum, 34, 51, 69, 78, 83, 112 note, 214, 216, 218, 220. Lymense, 69, 83, 102, 217, 218. numismale, 103, 217. Oppeli, 69, 70, 102 note, 103, 217, 221. oxynotum, 21, 40, 46, 69, 83, 101, 102, 199, 214, 21d; 216, 218) 221 Simpsoni, 69, 102, 217. Oxynotum Subseries, 106. Oxynotus Bed, 107, 109, 111, 129, 201, 213, 215. Zone, 106, 107, 116. Pachydiscus, 25. Packard, 3, 43, 47, 52. Paracme, x, 21, 107, 112. Parisian Basin, 87. Parallel Character of Differentials, 84. Parallel Series, 27. Parallelism, 22, 25, 27, 28, 43, 50, 53, 109. Parthenkirchen, 93. Pathological Species, 30. Pavlow, 115. Pelecypoda, 8. Peltoceras athleta, Pentacrinus Bed, 198. Pentacrinus tuberculatus, 153, 155. Perisphinctes, 24, 87. Perisphinctes aberrans, 33. Defranci, 24. Pern, 86, 87, 115, 118, 151, 153, 169. Petschora Land, 86. Pettos Stock, 23. Pfonsjoch, 122, 129, 134, 137. Pforen, 168, 171, 197. Phragmoceras perversum, 34. Phylembryo, 8, note. Phylloceras, 27, 122 note. Boblayi, 216. Phymatoceras enervatum, 24. Physical Selections, ix, 52. Physical Surroundings, action of, x, 52 note, 53. Physiological Equivalent, x. Pictet, 33. Pictetia, 24. Piette, 123. Piloceras, 12, 13, 52. Pinnacites, 51, 80. Placenticeras, 31, 32. Py 422 Soy Planorbide, 29, 30, 31, 78, 118. Planorbis Bed, 57, 85, 90, 91, 92, 94, 104, 110, 114, 115, 116, 122, 129, 134, 197. Planorbis Zone, 29, 90, 91, 92, 94, 104, 107, 108, 116. Planulati, 42. Plicatifera, 24, 42. Plicatus Stock, 25, 54, 55, 57, 80, 81, 83, 107, 112, 116, LON 22 25: Pliensbach, 210, 215. Polar Homozoic Band, 87. Poland, 86. Polygenesis, 118. Polyphylletic Origin of Forms, 31. Popanoceras, 4, 5, 24. Popanoceras antiquum, 4, 5, 23. Kingianum, 5, 128, 124. Portlock, 164. Portugal, 86. Pouriac, 115. Pourtales, 121. Primary Radicals, Primitive Radicals, 3, 16, 52. Primordialide, 51. Prodissoconch, 8, note. Progressive Characteristics, Genesis of, 71. Progressive Forms, Morphogical Equivalence in, 21, 22. Progressive Series, 25. Prolecanites, 3, 4, 5. Prolecanitide, 4. Pronorites, 4. Protection, effect of, 47. Protembryo, 8. Protoconch, 1, 8, 11, 13, 14, 49. Protozoa, 8. Provence, 91, 1138, 114, 116. Psiloceran Branch, 120. Psiloceran or Radical Stock, 54, 57, 71, 80, 81, 107. Psiloceras, 5-7, 13, 22, 28, 34, 85, 52, 54, 56-60, 71- 73, 75, 78, 80, 84, 85, 89, 90, 92, 98, 101, 102, 113, 114, 117, 118, 120, 125-128, 131, 136, 137, 155, 195, 197, 215. Psiloceras acutidorsale, 162. aphanoptychum, 125. Atanatense, 122. Berchta, 123. calcimontanum, 125, 124. caliphyllum, 54, 55, 57, 85, 90, 112, 117, 122. erebricinctum, 123. (goc.) ecrebrispirale, 124. (goc.) Portisi, 124. cryptogonium, 123. Hagenowi, Wih., 90, 93. 120, 123, 124, 174 note. Kammerkarense, 123, 124. longipontinum, Wiih., 91, 93, 122, 125. majus, 123. mesogenos, 21, 75, 124. megastoma, 96. 3.99 ><<. INDEX. Psiloceras Naumauni, 124. (Zgoc.) Naumanni, 124. pachydiscus, 123. planilaterale, 196. planorbe, 6, 22, 25, 37, 39, 41, 54-56, 58, 62, 65- 67, 78, 80-82, 85, 90-93, 112, 118, 115, 117, 121-124, 126, 136, 140, 142. 155, 156, 161, 163, 164, 195-197, 215. planorboides, 89, 122. pleurolissum, 75, 122. pleuronotum, 123, 124. polycyelum, 122. polyphyllum, 123. Rahana, 89. sublaqueum, 123. sublaqueum, Wah., 126, 147. tenerum, Wih., 126. Psiloceratites, 6, 72, 85, 89, 90, 93, 136. Pteronautilus, 80. Pteropoda, 8, 10, 16. Ptychites, 5, 6. Ptychoceras, 29. Pulchellia, 25. Pyrenean Basin, 87. Quadragonal Whorl, 56, 77, 78. Quedlinburg, 92, 137, 158, 146. Quenstedtioceras, 23, 28. Quenstedt, vii, 1, 29, 32, 33, 387, 55, 58, 62, 83, 90, 91, 92, 93, 96, 98, 101, 105, 112, 113, 121, 123, 125, 128, 129, 131, 135, 137, 138, 139, 141, 143, 149, 155, 157, 158, 163, 167, 168, 169, 171, 177, 179, 180, 181, 182, 183, 184, 185, 186, 187, 190, 192, 196, 197, 199, 200, 201, 209, 211, 216, 217, 218, 219, 221. Radical Stock, 22, 24, 57, 85, 107, 112, 120. Radicals, Acmie, 23. Geratologous, 28. Mesozoic, 22. Primary, 3. * Primitive, 3, 16. Retrogressive Forms, 28. Secondary, 3, 22, 71. Theory of, 21, 28. Transitional, 3. Raidwangen, 169. Ramert, 170. Raricostatus Bed, 103, 104, 106, 109, 146. Red Car, 96. Reineckia, 23. Replacement of characteristics, law of, 44; in anagene- sis, 74; in catagenesis, 78. Residual Basins, 89. Faunas, xi, 89, 106, 114, 116. Retrogressive Characters, genesis of, 74; tendency to inheritance of, 78. Retrogressive Forms, morphological equivalence in, 28, 30. bS Oo OX Reynés, 102, 103, 153, 167, 176, 181, 185, 200. Rhabdoceras, 28, 50. Rhacophyllites, 122 note. Rheetic, 28, 90, 103, 111. Rhine, 113, 114. Rhone, 87, $8, 91, 96, 98-101, 103, 106, 108, 111, 112, 188. ‘ Rhone Basin, 92, 96, 97, 102, 104, 106, 108, 111, 114— 116, 213, 217. Rinteln, 137. Robin Hood’s Bay, 146, 170, 171, 201, 203. Rollier, 90. Romer, 92. Rostrum, 29, 49. Rotiformis Zone, 109, 134, 189. Rudern, 121. Ruffy, 122. Rugby, 155. Ruminantia, 27. Russia, 87. Russian Province, 86. Ryder, vill, 43. Salins, 90, 144, 157, 169, 184, 197, 201, 204, 214. Sageceras, 4. Sandberger, Guido, vii, 2, 14, 18. San Francisco, 87, 175. Sannionites, 13, 48. Sadne et Loire, 87, 88. Saulieu, 91, 121, 142. Scaphites, 31, 32, 47. Scaphites umbilicus, 34. ventricosus, 32. Scaphopoda, 10, 16. Schaichhof, 191, 192. Scheppenstadt, 191, 193. Schlonbach, 92, 98, 101, 102. Schloenbachia tricarinatus, 24. Westphalicus, 24, Schlotheimia, 5, 6, 7, 40, 44, 54, 58, 71, 73, 75, 80, 84, 93, 106, 107, 113, 114, 115, 118, 123, 125-127, 134, 1385, 154, 157, 194, 201 note. Schlotheimia angulata, 98, 94, 126, 128, 129, 182, 134, 196. angulata, Wah., 180, 134. angulata, Zittel, 130. angulicostata, Gey., 135. aneustisulcata, Gey., 135. Boucaultiana, 94. Boucaultiana, Wih., 21, 75, 133. catenata, 7, 25, 40, 57, 71, 73, 75, 90, 94, 113, 117, 127, 128, 131, 134, 135. catenata, Wah., 25, 58, 129, 134. colubrata, 130. comptum, 134. Charmassei, 41, 94, 133, 134. Charmassei, D’Orb., 132. Charmassei, Wah., 152. donar, 128, 134. By A 2356 Schlotheimia D’Orbigniana, 94, 95, 133. extranodosa, 154. Geyeri, 135. lacunata, 135. Jacunatus, Gey., 135. Leigneletii, 94. Leigneletii, With., 152, 133. marmorea, 134. montana, Wih., 154. Moreana, 93, 128. pachygaster, 134. posttaurina, 154. Quenstedti, Gey., 135. rotunda, 135. scolioptycha, 154, striatissima, 129. Speziana, Can., 156. trapezoidale, 134. taurina, 154. ventricosa, 128, 133, 154. Schlotheimian Series, 57, 105, 106, 122. Schliiter, 92, 94, 97, 98, 101. Schwartz, 2, 14, 216. Scipionis Bed, 103, 176. Scotland, 88. Secondary Radical, Seebach, 94, 114. Selection, physical, ix, 52. Semper, 8, 30. Semur, 60, 91-93, 95-99, 102-105, 114, 121, 122, 129, 130, 132, 153, 155, 137-189, 141-146, 148, 156, 157, 159, 160, 162, 164, 166-169, 171, 175-179, 183, 184, 186-193, 195-198, 206, 208, 209, 211, 212, 216, 217. Senile Degeneration, 75. Senility in Orthoceratites, 34, 35. in Goniatitine, 35. less marked in ancient forms, 35. in the individual described by D’Orbigny, 36. Separation, Geographical, 27. Sepioids, 18, 26, 48, 43. Septum, 9. Shaler, 46. Sheaths of the Siphon (Endocones), 12. Siebenburgen, 111, 159. Silesites, 24. Silphologie, 9 note. Siphon, structure of, 11, 13. collar of, 17. changes in, during growth, 54. funnels of, 13. position of, 51, 52. importance in ancient groups, 51, 52. Siphonal cecum, 9, 18, 14, 19. Solenhofen, 51. Somerset, 92, 144. Sorbonne, 137, 138. South America, 86, 87. Southeastern France, 102. South England, 88. 3.99 Jere? 71. INDEX. Southern Alps, 87. South German Basin, 87, 94, 96, 97, 99, 101, 114, 118, 218. ; South Germany, 88-102, 104, 107, 108, 118, 114, 116, 117, 121, 135, 163. Sowerby, 68, 121, 158, 178, 188, 206, 219. Spain, 86, 108. Specialization, causes of, 52. Species, definition of, 56. Spezia, 116, 154, 166. Spitzbergen, 5 note, 86. Spheeroceras, 23, 32. Spheeroceras refractum, 32, 47. Spinifera, 23, 24, 42. Spirula, 26, 52. Stages of Growth and Decline, 14. Stephani, 115. Steinheim, 29, 30. Steinmann, 86. Stellaris Bed, 105, 109. Stephanoceras bullatum, 32. coronatum, 25. Geryilli, 32. microstomum, 32. nodosum, 23. platystomum, 32. refractum, 32, 47. Stephanoceras, Genetic Relations of, 32. Stephanoceratide, 23. Stonehouse, 215. Street, 92. Striaries Bed, 105, 203. Struggle for Existence, 27, 38, 50, 53. St. Thibault, 132, 135, 142, 144, 212, 213. Stur, 89, 109. Stuttgardt, 132, 156, 158, 176, 182. Suabia, 87, 89, 90, 118, 167, 180. Subclymenia, 25, 26, 80. Subnodosus, 188. 4 Suess, 90, 93, 95, 96, 97, 100, 109, 115. Surroundings, action of, x, 52 note, 53. Sutures, origin of digitations of, 50. degeneration of, 57. persistence in degeneration, 83. Sweden, 88, 115. Swedish Basin, 88. Swiss Alps, 114. Switzerland, 87, 90, 93, 114. Széklerland, 95, 110. Tenia, 45, 47. Taramelli, 115. Tate, 93, 117, 123. Temperate Homozoic Band, 87. Tentaculites, 10. Tertiary Radicals, 19, 22, 23, 107. Terqueum, 91, 94, 103, 117, 123. Teuchsloch, 203. Teutoberger Wald, 97, 98, 101. INDEX. Theory of Morphological Equivalence, viii, 21, 28. Theory of Radicals, 21, 28. Three Modes of Development, 37. Thysanura, 45. Thysanuriform larve, 9 note. Tilibichi, 86. Tingo, 153. Tirolites, 24, 124. Toulon, 114. Tmegoceras, 125. Tmegoceras latesulcatum, 125. levis, 125. Transitional Radicals, 3. 'Frachyceras aon, 22 note. Trigonoceratidee, 80. Trocholites, 35. Tropical Homozoic Band, 87. Tropites, 22 note, 24. Tropites Campigliensis, 154 note. discretus, 154 note. Jokeyli, 154 note. ligusticus, 154 note. subbullatus, 154 note. ultratriasicus, 157 note. Tuberculatus Bed, 102, 104, 105, 119, 144, 199, 219, 21d. Tuberculatus Horizon, 95, 98, 99, 100, 213. Tibingen, 94, 96, 101, 130, 132, 139, 143, 152, 158, 163, 179, 182, 191, 206, 214. Tubularia, 45. Turneri Bed, 159. Turrilites, 28, 33. Turrilites Boblayi, 31. Coynarti, 31. Valdani, 31. Typembryo, 8, 14, 18, 49, 52. Types, Cryptogenous, 117. Law of Eyolution in, 16 note, Tyrolites, 22 nole. Uhlig, 33 note. Unionide, 118. Unit of Classification, 53. United States, 86, 87. Uphill, 92. Upper Bucklandi Bed, 81, 95, 98-101, 103, 105, 107, 108, 139, 165, 188, 197, 206. Vaeck, 84. Vaihingen, 132, 143, 156, 157, 176. Vancouver’s Island, 86, 87, 118. Variation, 87, 89. Law of, x. Origin of, 48. See also Cope and Differentials. Veliger, 8, 14, 18. Vermetide, 115. Vermiceran Branch, 136. Series, 61, 112. 237 Vermiceras, 33, 34, 48, 54-57, 59, 60, 65, 70 note, 72, 74, 75, 81, 82, 84, 95, 96, 106, 107, 113, 114, 116, 136, 140, 146, 149, 151, 154, 156, 161, 162, 172. Vermiceras Bonnardi, 158. Bonnardi, Wright, 159. Carusense, 141. Conybeari, 21, 25, 61,72, 81, 95, 96, 115, 150, 155, 157, 159, 160, 161, 166, 168, 172, 175, 188, 194, 209. Hierlatzicum, 95. ophioides, 72, 78, 81, 87, 156, 160. spiratissimum, 57, 59, 61, 72, 81, 95, 96, 141, 147, 158, 159, 160. prespiratissimum, 96. Vienna, 86. Von Buch, 29, 32, 49. Von Jhering, 10. Von Rath, 115. Volcano, 172. Vortices in Eyolution, 108. Vosges, 118. Waagen, 4, 21, 27, 28, 33 note, 50 note, 51, 87, 88, 90. Wehneroceran Series, 57, 58, 79, 93. Weehneroceras, 7, 54, 57, 71, 80, 93, 118, 118, 120, 125- 127, 129, 131, 134. Weehneroceras anisophyllum, 127. circacostatum, 58, 127. eurviornatum, 57, 58, 127. diploptychum, 127. extracostatum, 57, 127. euptychum, 127. Emmrichi, 21, 127. (Aigoc.) Emmrichi, 127. Guidoni, 127. latimontanum, 127, 201. megastoma, 122, 127. Paltar, 126. Panzneri, 57, 127. tenerum, 126. stenoptychum, 127. subangulare, 126. Wahner, 6, 24, 57, 58, 60, 61, 85, 89, 90, 93, 95, 99, 108-113, 120, 122-124, 126-128, 134, 139, 140, 142, 147, 148, 153, 154, 213. Waldenburg, 188, 141, 144. Wallegg, 89. Waltzing, 157. Warwickshire, 92, 155. Watchet, 121, 157. Westphalia, 88. Western Alps, 114. Western Europe, 56, 60. Weissmann, 43, 44, 47, 49. Wellershausen, 142, 144. Whitby, 121, 146, 156, 166, 167, 169, 184, 197, 201, 204. Whiteaves, 87. : Whitney, viii. Wiesloch, 137. ere ct de 238 York, Museum of, viii. LS | Yorkshire, 88, 168. _ Wiltshire, 88. Winkler, 89. Woodward, 155. i | Wright, viii, 41, 59, 68, 92, 96, 98, 101, 102, 112, 113, | 121, 135, 146, 155, 158, 159, 163, 164, 178, 180- 184, 186, 188, 201, 206, 209, 211, 218, 217-219. Wiirtemburg, 123, 154. : Wiirtenberger, 2, 21, 24, 41, 42, 43, 61. Zeiten, 97, 149, 175, 188, 202, | Zittel, vii, 4, 7, 21, 82 note, 108, 113. | | Zone of Amm. Burgundiz, 91. Amm. Liasicus, 91. — ; Autocthones, 89, 96, 106, 114, 118. a (See also Tables 1.-V.) Xenodiscus, 3 note. | Ziirich, 34, 197. ¢ teienstedt, Fraas, Oppel a Cor. bisuleatum = multicostatum Sow. (not Quenst.) Arn. falcaries = falcaries (pars) Q. Cor. Sauzeanum Arn. kridioides = spinaries Q. = kridion (pars) = Buck. ecarinaries Q. | | Same — —Cor. kridion = kridion Ziet. (not Quenst.) q Means N09 engupnd oxlt shad en1990 - agony oe nay “inzdnafontes : sutonoliag Rite : TABLE I.—Genealogy of the Arietide in the Basin of South Germany, afteQuenstedt, Fraas, Oppel, and their Collections. NOTE. Q- = Quenstedt, and O,—= Oppel. ‘Oxyn. numismale =oxyn. “ (pars) Q. Oxyn. Oppeli WIE sxyni.numismalo (pars) Q Raricostatenbiy Cal. raricostatum Q Raricostatusmt i Oxyn. oxynotum Oxynotonlajt _ Sebi: lacunata 0 st | | Same Orynotusbat Schl. rotunda Tet, stellare obtusus suevicus Q. Ast. impendens ! Q Turnerifono, nb Ast. Broo! Ast. obtusum cae Obtusubett. cll. ces, | = capricostatus Ast, Turneri = obtusus (pirs) nudaries Q. Smithi Q 1 t Scipionis i} a pnianus olifex Q | diosa \ Agas, nodosaries Zvschonlagor, a) By Ast. Turneri E Bi Ce) or 1 | EDGE compressaries 1 Tworculatusbett \ Arn. cergs Turneri (pars) Turneri (pars) Q obtusus (pars) Q ‘Arn, tardecrescens =falcaries (pars) Arn. Bodleyi | | densicost: eraliteldes }(niiQ As slorat costa REnEE GIS Ast. acceleratum “ eet Cor. bisulcatum 1 | | =multicostatum Sow. Cor. trigonatum Zono of Amm: (not Quenst,) =nudaries Ieee railera Agas. striaries, = — — — Goometricus | | Same Arn. Same Arn. falearies Crossi Q. (not Wright) Ae Oe (para) Q Upr i Buckh Same =Bonnardi,O. = 3 = falcaries : if se | not D'Orb. C | ay | (pars) Q Cor. Gmuendense meh 0 = Cor. Sauzeanum = = ——— —_ — — — Sane Amn. kridioldes =spinaries Q : ‘ pe ot ata’ (para) Cor. orbiculatum SrtctoainarienO% Bucklandi (pars) mnecer i | “ —_ costaries (pars) a Cal. tl oe Caltaclentam i Am, ceras | ! oblongaries — ——Cal. ca © ———Cal. sule: Ss earatlten ~ | latisul. longicella Q. | erabltordes}( pars) }Q: \ | orllyra Cor. Bucklandi ° Arloteniageri 1+ Charma ! Same { Same nulticostatus (pars) eh s A ot plans 3, Fy previdorsalis = costal pars, Bucklandibett. s gigas. | pu a Froas i brevido 70 eunivtensis P Hower Buckland (pars) Q. Cal. Deffneri — Verm. Conybeari Arm, Hartmani | = solarium Q = c oe =faleuries | \ Cor. rotiforme——=Cor. Intum. = & Q Buck. pinguis? Q. =robustus Q. | zns. Imyigatum striaries (pars) Q Same | Jatisulentus Q Cal. longidomus Q Solil, Lelgnetetit =angl reaauis (pars) Q | Schl. Charmassei | anal compres (par) Q | 5 Angulatenbank, Schl Roan j = Arm. semleoatatuns — = 2h —-—---- — —Cor, iain | or angl: deprossus (pars) Q | : ate kridion Ziet Angulatusbott, } Sal eaves ee (not Quenst.) Schl, colubrata | angl Thittaaalens (pars) Q. Schl. atrintisimun Cal, carusense Taqueus i Q 7 Verm, spiratissimum | = Q striatus (para) Q Schl. eatenatn angl Phalaaslous (pars) Q. Psil. longi Inquonx fal. Dinssicum Gal. Inqueum Verm. spiratissimum! sironotus Q (pars) Q Taqucus (para) Q | 0. | Cal. John stoni Cal | | Inquens (pars) Q \ Upper part of this is tho Laquons or Cal ocuraa Dod Psilonotonbank, ) Sclil. catenata or ang, pallonotus Planorblsbett, * hireinus Q = pail. pli satus (pars) Q. eat Pail plat ro yar. plicatum =pailone | HQ Bonebed, Pail. planorbe var, love Gelbe sandstein, Soh), striatissimus f = psilonot. COG e} atrintus (pars) Q. ‘anid slbbiM } ‘ pingdnetsteoonsA P a ww 5 | _ StadeutsteooiisA ; : , mnpreantihtomtnitt et eg mal in t ae a Mitoe : ; HoxsinatonyxO, fu ' iat 40 . : Abavior Ads@ ; Re sttedeutonyxO Paes ; a | , | locttive nt 5, a4 anes] ia 2 Hadaiad : te | aan | { e Nap ws 6 nt ee TONEY CECA ¥ | ; Sane ® Fe neysinadoziwS wo SMedeutsluoisdu heey spearmint NB HYD eR Hen A are Arh, turdecréstans | iw Eotvaries: (pues) =densieosin ’ , Ba mamnmnA to atoS ifiseratile a : uy Lewantemosd | © ina ' “\haattontl r9qqU = bod | | ‘ ‘el eerie tyhG + ateemmnenet “pl Agi lcautu ir ake npcalle G 7 betes tinarhey segalaotaia iszeeortead’) ilo8 ath wh i ote) engiy sea) Coaaitn = Wiermne €2e dia oud etn) ibosblssa zsyod + i ; bad | Sanid | yi ek. fongidomya (, fAtipolentoe Q — Sa nn | : fitalonsio’ [dod ) (e1nq) anezorgaro! fynis = = ’ ‘eee snat 1D ita een Ayes = 5 ae Adadriotslu nA (ina) uhh taka “0 a » StadeutalugnA . At eeionstatim — poe duloa: Woe e (rag: eroieastad'D Sisiatan Ep ated oe rite Sp eniinei tes BIRD Io orm oepiras vit ”, paciiseua.\ (pars: it ‘ai Bilt 40 Nagg a7) ) he 10 auoripRD 9) ott ; -bod enr990.— fy, alisdnetonolie9 | ity Bienstno 1ilo8 sehaltars San = a te = Na card and Boucault’s Collection in Museum of Ast. Collenoti Ast. impendens um, R. Ast. Brooki Ast. acceleratum Ast. Turneri Ast. stellare Ast. obtusum = sits var. quadragonatum um | zeanum Cor. Gmuendense | — is = Isis R. | 2anum | A hele D’Orb. _ Cor. lyra Agas. striaries R. = Vercingetorix n part also = Terquemi R. ressaries : Quenst. Cor. rotiforme Same Agas. levigatum = Dall Ere R = Petri R. Cor. Bucklandi = sinemuriense D’Orb. = Ausoniensis R. Cor. rotiforme=——=Cor. latum = Hehli (pars) R. s====(or. kridion Cor. coronaries = Hehli (pars) R. = Hehli (pars) R. — Oor. kridion? — Agas. levigatum 1 ; ‘ 4 1 i i TABLE II.—Genealogy of the Arietide in the Basin of the Cote d’Or, after Collenot and the Collections per eds of the ) Tower Lins. 4 Bucklandi Zone. 4 Angulatus Zone. Planorbis Zone. Birchii Tuberculatus bed. =all the beds of the Lower Lias above the Upper Buck- landi bed. Bucklandi bed. = Upper Bucklandi bed. Scipionis bed. = Lower Bucklandi bed Angulatus bed. Liasicus bed. = Caloceras bed. Planorbis bed. Schl. striatissimum (bed not known.) Schl. Charmassei Cal. nodotianum Cal | | Schl. lacunata | | | Schl. Boucaulti- | ana | 1 | | J | | ! | ! | Schl. Leigneleti Same | | iS | | t | | | | { Same Same Jeet ) j jl ! | | | Schl. D’Orbigniana il es | | | Schl. angulata Schl.colubrata Psil, longiponti- num Schl. catenata | raricostatum ! | | Cal. Liasicum —$<—>_==>>$_——=Cal, Inqueum = Burgundia Mart Same at Semtr, and Réynés, and Coll. = Collenott. Cor, bisuleatum, D'Orb etR tenuarius R. Same ' Cor. bisuleatum yar. like Sauz um Arn. Hartmanni Cor. Sauzeanum auubrius R =nodosus D/Orb, Orioceras Fryon R. = Gaudryi R. and named in par Arnioceras in all forms as compress including Arn. mise. Quenst rabile and Arn. kridioi des. (See Table V-) Cor. kridion——Cor. = Hehli (pars) R Arn. falcaries —--—- — — — — — — Cor. = Hettangensis (pars) R. ridion? Boucault’s Collection Ast. Collenoti c aie: Ast. Brooki Ast, acceleratum Ast. Turneri Ast. stellarc———— Ast. obtusum. = oR, var, quadragonatum Same Same 1 | Cor. Gmuendense | Isis R | Poel Cor, Lyra Agas. striaries ercingetorix rqnemi R t Cor. rotiforme Same Agas. lavigatum = Dall Ere R = Petri R Jandi nse D'Orb. Cor. rotiforme———=Cor. latum = Hehli (pars) R. ‘oronaries Hebli (pars) R. Agas. evigatum in Museum of Comparative Zodlogy, Oxyn. Lotharingum = R Oxyn. Guibali = ars) Juibalianus (pars) | K reenoughi Manus (pars) R = Guibali (pars) R Agas. Scipionianum——Agas. Scipionis war aries Reson r ‘hinder eC = Fete ' ; Ean pire), | Sy Pre: euteluareduT \ 9H) to shod i > hob ite tay es s#hid Ta8v0 z Lariat H..* - a siaoupal Jiio2 -‘Hlpesuedl los oe evods exid towel -Hoyl tsqqU ort _ bad ifnal ay E y Th $ ' vay : " | non i iowa ert i eels’? ean. ag Ahora: Same » SAN > I Halsagiat [doe | Varo TOBA wewmee Virgiy Or thin : = Ayer eth ef Comnourantee RR , * bad ibnaitous ; . | j ibasldoud taqql = a E | i had f .on10N ibastdoutt a ie i m~ " $< sepprechaesles seabed AS SN | Oar i Vern, ephicides rit | 5 ‘ beet ainoigioe ey ae } { ibasidovd rod = : * : cone " yeh stat benct “hodel. caThisetas Mesiane = Rodsjia 0 2) ie Me fate ft ~ -bed autrlugna welll " 24 tee ri ome Fie ony bear? ieee ire: | pees} tyeiag. ‘ Giatiinclianie, loa ‘ (ftwont ton bed) -bsd euoiesii -bed eptoc0laD = ; ' - Dumortier. Middle Lias. Oxyn. Oppeli? = Oppeli D. | Planicosta bed. = Raricostatus bec il tis D. ] Oxyn. Buvigneri = Buvigneri D. I Oxyn. Guibali Oxyn. Lymense = victoris D. = Saemanni DD. Oxynotus bed. = sf bed. Oxyn. Simpsoni Oxyn. Aballoense = Oxynotum (pars) = @ D. Oxynotus Zone. 4 Ds Oxyn. Oxynotum = Oxynotum (pars) D. | Oxyn. Guibalianum? “ 1D), Stellaris bed. a = Obtusus and Tu berculatus beds — —Agas. striaries = Davidsoni = Berardi (pars) D. Davidsoni bed. 1 (Striaries bed). Agas. levigatum = Upper Buckland = Berardi (pars) D. bed. - TL Buckland Agas. striaries - — — Agas. Scipionianum Bucklandi Zone. cc = Davidsoni D = D. Angulatus Zone. = Angulatus bed. | Agas. eyesem 1 See Part II., p. 174. 2 This is a form mentioned in Part I., p. 115, probably a species of Liasicus bed. Coroniceras, allied to Cor. kridion or rotiforme. = Caloceras bed. 3 The species of the Arnioceran series are arranged in line accord- ing to the beds in which they occur. It was not possible to trace the genealogy accurately. ' Planorbis Zone. EO me et te a ee ae ane Pc. ‘ 4 | | ® tasoispal tae} adiongly fiat - Ch atbanpanid - At ae ah oe eee zodols 1K: pares 2b Saah cr regis eA ME TABLE TIT. — Genealogy of the Arietida in the Basin of the Rhone, after Dumortier. NOT: = Dumortier. Middle Lias. Oxyn. Oppelit = Oppeli D. Cal. raricostatam = Pellati by Arn. Macdonelli armentulis =noilotinnus D. Cal. carusense vellicatus D. = Vdmundi D. Amm. tardecrescens D. of this formation is Arnioceras (sp.!) not determinable Planicosta bed. = Oosteri D. Raricostatus bed. Ast. Collenoti = Cluniacensis D. Am, miserabile =Jejunus D. Oxyn, Buvigneri ia D. Oxyn. Guibali Oxyn. Lymense = victoris D. Ssemanni D. Arn, Bodleyi? | Oxynotus bed. = "bed. D'Orb, & D Oxyn. Aballoense = Oxynotum (pars) o D. ; Dd ynotus Zone. 4 Oxyn, Oxynotum = Oxynotim (pars) D. ‘icostatum ! WB eet Oxyn. Guibalianum? al. earusense Arn. Bodleyi aa Stellaris bed. ST =Tandrioti D geometricus D. =Obtusus and Tu- } Sell. Boucaultinn | Ast, stellare ulatus beds \ = stellaris D. ! Ast. obtusum | RED: 1 Schl. lacunata Inchinatus D. | Cor. bisuleatum. resurgens D. — — — ~Agas. striaries i Davidsoni | Cor. bisuleatum Berardi (pars) D multicostatum 1. ' Parideonl ped Same! Ver. Conybeari Arn. kridioides? ! Cor. kridion Agas. levigatum (Striarie Buckle ‘i “1D. debilitatus Rey = Patti D. Cor. bisulcatum, = Sauzeanus 1). = Berardi (pars) D. SUPpeg Eckard Falsani (pars) D. bed Arnouldi D. ! micostatum — Cor. Sauzeanuy——— Hartmanni D. (para) D. Or, Gmuenlense D Ver, Conyheari? metricus 1. 1 | | D. 1 Cor, Lyra Cor Bucklandi Agas. strinries - ~ — Agas. Scipionlanum , =Lower Bucklandi | Schl, Charmassei = Arnioveras (sp.1) = wureus bisulcatus D. =Darvidsoni D ) Buoklandi Zone, + hed. (te aS) Arnouldi D. =hisulcatus D (pl. 2, not pl. 8) (Arn. Hartmanni?) * (pl Ver, spiratissimum? | = ‘ D. | ———$———Cor, rot Unknown form hisuleates (pars) D. y Schl. catenatum - — Cor. kridion Agos. levigatum Angulatus Zone. = Angulatus bed, ORaIATHEND) “op. D Cul, Johnstonim——Peil, plunorbe Cnt. lagu ' See Part IL, p. 174 = Dd! D Burgund 2 This is x form mentioned in Part L, p. 116, probably a species of | 2 , Coroniceras, allied to Cor. kridion or rotiforme i Liasicus bed, Planorbis Zone. = Galacérnai Led 1 ‘The species of the Arnioceran series nre arranged in line accord = = == = ing to the beds in which they occur, Tt wax not possible to tmce the genealogy accurately A mm f ~ wmateovtiy In) AMD yA * “ Hallol = Labs : 5 silniganris = E : ganoeune) La * plogisiv == , ap oad , A ibauatba = Cf atuvilley = raat wiaotl stad vim tet i crugpol, \ — \ ———— | ieee | a oe ats nod yuo") a fot (7A Siyraatod = Ts dvr sine Cinitasteooires TaD ry “4 Ss ssergaiyias ta) | k ijoinbant = s pbs orth vine lao both cass (SiT1OSy = «bad 2 ye? foe ane Ty) abed erctela } I 6 5 ; ‘ ginmryat (foe {. ‘eben ae f | AT artantiont - ! ; Avett aii th irae 10D 734 Sgntaert) 1) | = é ty Tita’) 2 — tydi sutaiilideh = Sse: ; cig , ‘ [oe pede ' ae he ee ; ie I Se’ 33 igs avy fy TEMES cope nner mem ¢ ay onquntl Aaryd 2, { ae ; eet ote v ie iu tas pow = Sirnstivnol? ta | \ qe gf THOUS. | Toness = j 3 erro ricats | {yi h Pie Cay ral * Say Picnate His a rrhikentig I / pai a mais Ae) iPetal fave xe SheMale hye. oe it # = F Nie ra) eho | -§ ye ae Pte + 2 { Soninpigemene? 19% et Nae hd H H at ue - pi ee SAO NY ee Maes ae Npiehal Se. : % = Se a of ee a eters ‘3 inndnagyay foe. ; ee ; ; | AE autuiegek = ; f bedded lie} 2 ie omicanih Lieder tore nitOh) int} ALG | sniboersaneefl-— <4 figs "- SOS a ce Oe pete Neer : ; ; ¥ “ TABLE I\d various Collections. } | 6 Oxyn. memismale Miele ies: = Amal. Wiltshirei | W.. Cal. aplanatum Raricostatus Zone. = Ariet. tardecrese¢ — CC bed. Oxyn. Guibalii I Oxyn. Lymense =Amal. “ W. Oxyn. Greenoughi : ] = Amal ee Saynetis eon peat OTE Ww | obtusum Oxyn. Simpsoni | Tet. =“ =Aymell © goc. sagittarium | s Slatteri Oxyn. oxynotum Amal. Guibalianum (pars) W. = Auman Ww. | I | i I Obtusus Zone. Seeds Agas. levigatum — — — — — PATI Sa Sow. Turneri Zone. j | = Tuberculatus bed. 1 | — — — — — — —Agas. Scipionianum = Ariet. ss if Upper | Bucklandi Zone. 4 Schl. Boucaultiana ze 7 bed. == ss We | | | | -Lower | Bucklandi Zone. 4 Schl. Charmassei = ee bed. —igoe, “ W. | Schl. angulata =—Z roc NS Wis Angulatus Zone. | = ue bed. Lower part of this zone = Caloceras bed. Schl. colubrata = Mgoc. morianum W. Schl. catenata = Mgoc. “ W. Cal. Johnstoni =Mgoc. “ Wi | 1 May al - , Bas earn mrclcner (ears) ay also occur in Oxynotus Zone, according to Wright Planorbis Zone. | I | 2 See Wright’s Plates, pl. 6, figs. 2 and 3. 5 bed. 1 Psil. planorbe SS ® The lines connecting the species of Arnioceras indicate = ae @ genetic bonds only in a very general way. The true succes- = “ erugatus sion of the forms is not given in this part of this table. Hanobow lt nh. Iwo .pgomA = + yitataoviner nD i maaan: irae W eunsiobon J9ith = WY pS eB ES Went tge dsith = aangeute kD ae eee = ode - —— ba Pi rr FE “wlanuoal Loa WW Thee nope <= , sue amae cma 9¥ | nee a ‘ bod a W ensintornosy = Wiliheokh Soins Ae eat. F 5 end add anndis rie’ i iyalhind wins» "Gi atearroltib: soit = po mateseosiiaes Winaq) | 4 innaminehl oil apo) aA ve ~ pnettlunoiiotl Aae eae at aireoxotostrist JntA ( : { (dangle ction a1 ROOT N82 ai 3 WW (zing). ®toi wiyokharwd ww J _ Pio eiayrydi ‘ Salers Ayre ipa WY innadigaoD sion irienne ; Be re oraed engra) ie Be Ses JaitA = = B tl Dinoe Ss mis 4 “af Tt Nurisell Siige"t9 Winn ? Fe t (nipinooay n6iitear ) ee En Gl ma AL itned unio) Agitd © * msupet fe umoianil a0) “W auilosupal Soy. = cou pal | fig aE Pe (ennq) inwibsevieiai sogth = oa TABLE IV.— Genealogy of the Arietida in the Basin of England, after Wright NOTE, — W. = Wright, and B. = Blake. Middle Lias. Arn, Macdoneli al, raricostatum =Amm. “Port! Chl, nplanatum = Ariet. nodotianus W Raricostatus Zone. Ariet. tardecresven; Aric. “OW. z Bed | % i Cal. carnsense ll | Ast. Collenotr | = Egoc. Slatteri | (pars) W Ast. denotatum: Oxynotus Ait Sel LEC - - nilenoti (pars) W Ast. impendens = Ariet Collenoti (pars) W. Ast. stellare Ariet, * W, Same Same! | Obtusus Zone. Ast. obtusun, and various Collections. Ast. obtusum Ariet. Zgoc. sagittarium Slntteri (pars) W. Agas. levigntum — bed ‘ Ariet. * W Amm Amin, Sm Ast. Brooki Ariet. " W Ver. Conybeari 5 | = Ariet. Bonardi W. geometr Ast. Turneri ———- Ast, obtusum =Ariet. “ W I Turneri Zone. TVuberclatus bed Cor. Gmuendense Ariet. Crossi W Bodley Cor. bisuleatum et. difformis B. = Ariet. subnodosus semicostatum (posit. uncertain) W. 1 (pars) W. Arn. Hartmanni Upper | 1 Bucklandi Zone. } Schl, Boucaultiana Arm, Gerns bed * W 1 Arn. tardecrescens | Arn. semicostatuni———Arn. falcaries Cor. Sauzennum Ariet (pars) W Ariet. “ W. i | Cor, lyra = Ariet. bisulcatus W. Cor. Buckles = Ariet 1 Lower Cor. rotiformne———Cor. Bucklandi Bucklandi Zone. Schl Charmassei Ver. Conybeari = Ariet. " (pars) W Ariet. sinemuriense al = Fen W Ariet. "W. B | Ver. spiratissimum (position uncertain) Ariet. Gonybenri It Schl. angulata ! Egoc. “ W. Cal. Inquenm Angulatus/Zone: SehL colubratn Tnyrergnt of iazone 4 = 40%: morianun W Galoceras bed. Cal. liasicum Hoe. laqueolus W Same ————Cal. Inqueum Egoc. intermedinm ( Belclieri (yiirs), W. Arm. intermedium Portl Cal, Jolinsto Cal. tortile Egoc. “ W. ZEgoc. intermedium Belcheri (pars) W (pars) W. Planorbis Zone. bed (Authors) Ego. "* W. Amm: Jobnsto =A) s Sow =“ erugatus Bean, ! I 1 Psil, phinorbe var. eye—_—$—$—$— — Pil, plianorbe yaar. ppbie et ta my) —Agas. Scipioninnum ‘ ‘The lines connecting the spe genetic bonds only in a very general way Oxyn. memismale Amal. Wiltshirei 1 W. Oxyn. Gnibalit Oxyn. Lymense ‘sat "OW. Oxyn, Gr Oxyn. Simpsoni | Amal. “ Ww i) Oxyn. oxynotum Amal, Guibalianum =Amal. “ W i W. W 1 May also occur in Oxynotus Zone, according to Wright 2 See Wright’s Plates, pl. 6, figs. 2 and 3, 8 of Arnioceras indicate The true succes sion of the forms is not given in this part of this table. ee cou tana ere ( Bi biked _ dine as soonnt KO muset Ee) f itolomgial filoe eee msc rt susilosgoo8t lito Teed Nelnatt is Deh rds iva: : bed + xadl wbed inozemsL Vee itenAiiectacs oe oat aon” metthare.s 7 eee i : hddaprrmeniee ‘bed 2ufonyxO “bad zueutdO~ By N euentdO 5 hata pe ray a 4 sbed eutsluoveduT 4 ; Tiere tne tvaud ie 09D) Aw. sbed 2uoiits ibnsbiova 1gwol so smn omiae Solistot [nO (baslyot) teecoee : omse en Aieh ae ABD) inote fuori l In a ll omen ; . ee iE he fearing | ofittod [nD oes muNnoilg KT ,sdtosely at iin dh Mead on Seba nehechs ane nee e ebnei elbeiiiei ete pilin cee Tt nemeetae coe =a iE vat or ane .bed chads t ‘taloagia ‘toe sae hemere er Mk pine iosencrial toe ar auings EE etalngie [oe SMAirn. jiiechahsiy steiduloo [ioe ‘bed'zgye001s0° athena) Gar eee Poe 5nd a fe F X, .bed 2idionst t {onsen eve. FPF a ey ine ~—{oldanuiteanpy antigetininie . (nisadl mirigy oF) i Nahner, Mojsisovics, Cal. suleatum H Amm. Nodotian Cal. carusense F Amm. Liasicus | Cal. doricus Hy. Ar. “Gey Cal. Haueri Hy. Ar. sp. ind. Gey( Cal. Haueri? Hy Amm. “ Giil Amm. euceras ‘ Cal. suleatum Hy Amm. Nodotianu Cal. salinarium Hy Cal. carusense H Am )5* Haner Amm. epiatseiuy Amm. Liasicus Cal. Grunowi Hy. Hy. Ar. Gs Wih. Wah Cal. centauroides Hy. riHy. Ar. ne Wah. | Wah. Cal. prespiratissin 4 c Ar. “ | ih. Cal. latecarinatum Hy. | Ar. < Wah. m Hy. Wah. | ————————, Ol. Sebanum Hy. fig. ee Neum. MIDDLE LIAS. LOWER LIAS. THE FAUNAS OF ARIETID PARACMATIC FAUNAS ARIETIDE, ACMATIC 1 ARIETID.E THE EPACMATIC FAUNAS OF Oxynotus Obtusus Zone. Bucklandi Zone. { Angulatus Zone. Planorbis Zone. { Rhuetic Zone. TABLE V.— Genealogy of the Arietide in the Province of Central Europe. -_-_ Henleyi bed. Ibex bed. se Jamesoni bed. Oxyn. numismale Same Oxyn. Oppeli Same Oxyn. Lotharingum Oxyn Lymense Oxyn, Buyigueri Oxyn. Simpsoni Oxyn. Gaibali Oxyn, Aballoense Oxyn. oxynotum Cal, aplanatum Raricostatus bed. 4 1 Arn. jejunum Same? Arn, Odsteri Ast. Collenoti y ' ; i Arn. Macdonelli { Same Same Same Ast. denotatum Schl. Same Same? 1 Oxynotus bed. 4 Sdn Same Same | Gal. rnricos- | aa SS tatum @AREEh 1 Same | | Ast. impendens———Same Same Same Same \ | Same? | | i | Tuberculatus bed. Same Same gas. nodosaries Same Same Ast. Brooki Ast. Turneri ——=Sn Same Same i} 1 1 U ! — | Schl. Boucault | Bodeyi Schl. lacunata | | ela | Cor. trigonatum Cor. Gmuendense Wears J Schl. Leigneleti —--| a | a Same Same Same Same Same ] Same Cor, orbieu: Ast. stellare—=Ast. acode- | Gal. Nodotianum —$—__—__—$_$___——Same |e he latum | Ast. quadmgo: ratum Geometricus\bed: Same | 1 acum | Same ! Same Same | Ast. obtusum Same Same Same 1 1 U 1 1 i Cor. Bucklandi | Cor. rotiforme————————Cor. latum | Agas. Scipionis ! Same————Verm. ophioides i} Lower Bucklandi | Same —_Agas. stri—Agas, Scipionianum: bed. | | Arn. tarde- Arn. cerns Jartmanni Same Arn. kridioides Cor. bisul- ---Cor, Sauze-——Sime ——————= Cor. coronaries nries 5 Cal. Deffneri crescens | | catum anunt | Same 1 Cal. tortile?’ Same Same | \ (England) Cal. longi. | | | domuiny | Same Arn. obtusiforme Same —----— SA ——— | ! ! T Schl. Leigneleti | | | | | Gal. Inqneoides 7 | | | 5 | Y Schil. Charmas Cal. snleatum ——| ‘Arn. falearies Cor. kridion Agas. levigatum. Angulatus bed. 4 \ = 1 | | i I | “ Cal, carusense=___Cal. Jaqueum=—=Verm. spiratissim= Verm. Conybeari Arn. miserable? - Se ——— Schl. angulata | mum | tatum | I Same — — — Schl. colubrata | | | ~ | Cal. Linssicum ' ! | Galoceras bed. 7 5 y Same PailjJongipon- (Same Same———=___Cal. Inqueum ~-— Verm. spiratissimum? Pail. Hagenowi | num. | | | Schl. catenata Cal. John- Cal. tortile stoni Planorbis bed 4 Same 1__pyil. planorbe, var. plicatu¢—<==_<_$_<_<_$_$—_— 7 “ | ane —— planorbe, va plic | $$$ III 1 | i i i i ee Bone bed. Psiloceras planorbe, yar. leve (So. German basin) ae ee eee 5 1 Schil. striatissima (questionable) Gelbesandstein. (So. German Basin) the Lower Bucklandi bed. Oxyn. Greenoughit 1 This occurs, according to Quenstedt, in 1998) atuolueiddngin lye IL ita you .oA Taye) stannonl doe atl alatecoinnat oe dol yok — AVE isregperiad’) .o2 mp !-*" cometh = GW aeooiunoy Tie doe EW neovittnay vin? HEV dyin ufos ne iobins) des 2 Sisraacath alo Wi © animetnormitnl ie ¥t au me a mudoxtqorate Bits i * muieyique doW “Ok cnullydqoaian lon » quttloyiqolqail tes W » mutantoivaua ite ie mitaseooneti lag M: cheat ea eR * p prmahnth alot oh ae sat tat at ge i co en a * pobmatieoy toe é rat oil oyiqeilone Ho aw YF atilugan oe B. hott Pe ecanca ABT ‘ MEW? Bein Bitch ss (e1%q) ane 9, si 97 at iat Wi atetdulos aly? PueSTONn ane nouae ~ aotontan toe : AVE a 19328 ion toe We : . b salen jd ioe ra + Set a digi aw sna0tl ab? sehllaad ae fi Gone: ‘itonbA. Nstdoide@-sialiotH | (sgiomnosoel4 oF ibnaldond aq = bad eointeoormndl 3 At Pee atte oe bed 2inictiioA ‘bad ibaal dont ne nO = at es eae ‘brs sevormaM ie eat e— EXPLANATION OF PLATES. ALL specimens not otherwise described are in the collection of the Museum of Comparative Zodlogy, and all not mentioned as “casts” have the shell present, either in part or as a whole. Paul Roetter drew the figures in outline by measurement; the author redrew all the specimens using these outlines, but testing the accuracy of the measurements before they were finally placed on stone. The outlines in the Summary Plates were sketched by the author, and redrawn by Miss Pierson. Unless otherwise specified, the figures are approximately of natural size, although the process of reducing by photography from the enlarged drawings has introduced some slight deviations from the measured diameters of the originals. TABLE VI. — Genealogy of the Arietide in the Mediterranean Province, after Hauer, Neumayr, Wahner, Mojsisovics, Herbich, Giimbel, Geyer, and Rothpletz. ——— nat Arn. Bodleyi? Hy. « Amm. © — Giimb { Arn. falcaries? Hy. Amm. “ —— Giimb. { Arn. ceras Hy Amm. “ Hauer Am. tardecrescens Hy Amm. Hauer { Arn. semileve Hy Amm. “ Hauer Cor. Gmuendense? Hy Ar. u! Geyer Ar. ae Rothp. Ast. Collenoti Hy. . Adneth-Schichten, Amm. difformis (pars) Hauer Oxy. “Geyer Hiorlatz-Schichten, r . Geyer Fleckonmorgo } Arn, Suessi Hy a Hauer = Upper Bucklandi to 5 Cal. suleatum Hy. Pail Geyer Cor. young : y c4 bed Sch, angustisuleata Geyer Amm. Nodotianus Hauer Amm. “Hauer Ar. sp. ind. Ge Oxy. Greenonghi Hy. Raricostatus . , ly ‘ = pl Aa Brooki? ae Anmm. hy Hauer i Cal. carusense Hy Jor. Hungari mM ‘othp Sch. Geyeri Hy Amm. Liasicus Hauer Arn. abnorme Hy. Amm. Oxy. Guibalianum Geyer Teaeets Pal. “ Geye Am, oxynotus (pars) Hauer Sch, lacunata Geyer Cal. doricus Hy. Ast. stellare H Ar! yer Arn. cuneiforme Hy. Cor. bisulcatum? Hy. an yer Oxy. Lymense Hy , Ar. Quenstedi ( Awnm. “Gib, Anim. oxynotus (pars) Hauer : Cal. Haueri Hy. dwarf Verm. Hierlatzicum Hy. Ar. ambiguus ch. tenulcostnta Hy Ar, sp. ind. Geyer pl. iii. Amm. “Hauer. Ar. ampliceres x. Wa fig. 16 Ar “Geyer. Cor. Bucklandi Hy. Agas. levigatum Hy. Ast. obtusum Hy. Oxy. oxynotum Hy. Cal. Haveri? Hy ar. sinemuriense? Amm. abnormis Hauer Amm.stellarisHauer Oxy. “Geyer eye ay iNaeh, w (Chaen Avatiiserabiletiiy, AmmsBucklandi Gimb, Cym\ globosus|Gever: eit ivarnaaeciliis =e ver ‘Amm, cuceras Amn « Amm. laxvigatus Giimb. Sch. Charmassei Hy. =Amm, "Hauer III | Sch. aff. ventricora With Cor. bisuleatum Hy Amm, multicostatus Hauer ae J Cal. suleatum Hy Verm. Conybeari Hy Enzesfelder-Kalk, PCAN AU AmimaNcuoliniu caver A icc ae ater Cor. rtiforme Hy Rotiformis bed, aaa Sohincollcnty eli Sch. ventricosa Wah eateries tani Cal. ophioides Hy Amm. “ Tauer Bear Buckland bea, } Atm. mon fre Rei Ry Se ea teee leat Aye Are ue Walk Cal. salinarium: Hy | Amm. Hajcr Amu. spiratissimus Hauer Cal. Inqueum var. Scylla Hy. Amm. Linsicus Hauer Ary pet “ Wah. Cor. kridion Hy I Amm. “Hauer a Web. Emmrichi Wih. sp? | \ Weel, Guidoni ies | st. stellforme Hy Ar. co With. Sch, marmoren Wwh. diploptyehum “ # | Wiih Sclh, trapezoidale Cal, abnormilobatum Hy With, Wet, Iatimontanum “© { Ar. “ ‘il, | , Gor, kridion? Hy: We. stenoptychum “ | Amin ca Moja. Sch, pachygastor | Cal. Grunowi Hy Wall. Web, cuptychum = | Cal. Castagnolai Hy Ar Wi. | | Wer wal = Ar. Wiili | mh. anisophyllum " | Cal. supraspiratum Hy. Macmoren andi | 1 Pail. Kammrkarense Wall Ar. Wah Cal. centauroides Hy SES Rat, 4 Wel. haploptychum | Sal. Detzkirchneri Hy. Ar. Wah eer | s bi Cal. nigromontanum Hy Cal. perspiratum Hy. Ar. Wah aloce : We curviornatum “ “ Pail. caleimptanum —“ | = Hx. uy Wih Ar u Wah 1 Cal. prespiratissimum Hy. = ! anal , | 1 Cal. cycloides Hy. Ar. « Wii, Sch, angolata Wah. Werh, cireacostatum “ " Peil. pleuroptum =“ | unnanied Cal. fig. 6, pl. xvii Gal. Coregonense Hy. =Ar. “Wilh Cal. latecarinatum Hy Soli, extranodosn var. undetermined.) | Wii Is i Wah 1 i " Wal Wah Wich. Panzeri “Pail aphanotychum —“ | Seebachi Hy I Cal. gonioptychum Hy i | ae Wih. Cal. Haueri Hy =a Wii, d iy Woh. Frigga Psil, Bercht © | Psi. mesogenos Wal, Cal. Hadroptychum Hy. Arn Neum Sch, Donar Wah, Sch, taurina Wal). | | With. Cal. Loki Hy «Hy, Pail. crebricinctum Wah. Psil. Atanatense Ar. Wil. Num | Weel, Ralhann «4 Pail volyptatum “ 1 I Sell. montana Wal. Psil pleurolissum “ Cal, Juhnstoni Hy Cal. Linsicum Hy. @ optychum H. ! Wah Patiar «= Peil. pachy\scus “ Psil. sublaqueum With ; 1 dEg, Wii yr. y Sea tea AN Psil. polyeyclum \ re Sch. angulata ----- Peril. Gernese Wah. Cal. Johnstoni Hy Cal. Sebanum Hy eal xa a Seti Slt, catenintn Wath, Pail. majusyah, pe eee potdes rah: _ 1 This variety ia fgured in Mojsis et Neum, Beitr, IV. pl. xx. fig: 6, and Planorbis bed. 2g, angulatum (para) Neum, is, we think, 1 distinct species = Aig. subangulare Neum, ot V Pail eryplogonium Wal Beil. Hagenowi Wah bi List vs Wiehnerocerns was so long that ¥ could not use customary nomen- deg. uh eum. Amm, jun! clature, and liave referred each species t0 the describer of the specie: e ° Hy. scriber ¢ pecies instead of i mise Byatt the describer of the genus ul Pail. caliphyllum With. Neum PLATE I. Psil. planorbe. Fig. 1, cast with incomplete living chamber, showing folds at an early stage of growth. Fig. 2, suture much enlarged. Loc. Whitby. Fig. 3, 4, cast with similar folds in the young, but smooth in the adult. Fig. 4a, young of Fig. 4 enlarged. Loc. Neuffen. Fig. 5, 6, cast of the more involute variety, loc. Balingen. ‘The specimen has distorted sutures, showing the broad abdominal lobe and the large median saddle on one side, but is otherwise normally formed. Cal. Nodotianum. Fig. 7,1 specimen reduced to less than one half, showing incomplete living chamber. Fig. 8, sutures enlarged. Fig. 9, a fragment of a cast showing sutures and part of living chamber. Fig. 10, section of four whorls. Fig. 11, a cast. Fig. 11 a, section of last whorl. Loe. Semur. Cal. tortile. Fig. 12, specimen with portion of living chamber. Fig. 13, same, portions of the two outer whorls removed, showing their rounder outlines. Fig. 14, suture of same, enlarged. Loc. Semur. Cal, carusense.? Fig. 15, cast of the younger stages, broken out of the interior of an adult specimen, loc. Balingen. Fig. 16, cast, loc. St. Thibault. Verm. spiratissimum. Fig. 17, cast, living chamber incomplete, loc. Nellingen. Fig. 18, specimen with distinct channels, developed at an early age, loc. Semur. Cal. suleatum. Fig. 19, specimen with incomplete living chamber, showing the smooth young in the centre. Fig. 20, suture of the same, enlarged. Loc. Semur. Verm. ophioides. Fig. 21, cast of a broken specimen, showing the well developed chan- nels and keel in the young, and the early appearance of the pile. Figs. 22, 23, sutures of this and an older specimen, enlarged. Loe. Semur. Cal. raricostatum. Fig. 24, specimen, showing the senile metamorphoses. Tig. 25, section of last two whorls, showing the corresponding change of form for comparison with Fig. 25a, the adult of the same variety.* Loc. Balingen. 1 This figure is not numbered. 2 See Plate II. Fig. 1. 3 See Plate VI. Fig. 15, for the Jofnstoni-like variety. PLATE |. GENESIS OF THE ARIETIDAE. HELIOGRAPH, HART & VON ARN, N. Y. ROETTER & HYATT, DEL. PLATE II. Cal. carusense.! Fig. 1, specimen reduced to one third, showing incomplete living cham- ber and old age. Fig. 2, the same broken so as to show sections of the old whorls and the depressed abdomen of the adult stage. Fig. 3, enlarged adult sutures, also shown on Fig. 1. Fig. 3a, same, but very old, also shown on Fig. 1. Loc. Semur. Arn. miserabile. Fig. 4, large smooth specimen, with sutures and incomplete living chamber. Fig. 5, cast with sharp abdomen and gibbous sides, living chamber incomplete. Fig. 6, enlarged sutures of the same. Fig. 7, var. cunetforme, with incomplete living chamber. Loe. Semur. Arn. obtusiforme. Fig. 8, section of a cast, showing form of internal whorls. Fig. 9, specimen with incomplete living chamber. Fig. 9a, enlarged sutures of fig. 9. Loc. Semur. Arn. semicostatum. Fig. 10, variety with incomplete living chamber, the young remain- ing’ smooth until a late period of growth and ribs immature, loc. Semur. Fig. 11, cast, with ribs earlier developed than in Fig. 10, and slightly more prominent. Fig. 12, young from same blocks of limestone. Fig. 13 a, sutures from other young specimens on the same block, of different ages, all natural size. Loc. Whitby. Fig. 14, cast with folds in the extreme young stage and true pile beginning afterwards ; otherwise the form is perfectly normal, loc. Basle. Fig. 15, normal variety with deep channels. Fig. 15a, suture of the same, enlarged. Fig. 16, specimen with incomplete living chamber, channels developed at an earlier age than in normal variety, and abdomen broader and flatter in proportion than is usual in the species at any age. Loe. Semur. Arn. Hartmanni. Fig. 17, cast, loc. Bonnert. Fig. 18, suture of the same, enlarged. Arn. tardecrescens. Fig. 19, cast of a young specimen with incomplete living chamber. Fig. 19a, enlarged suture. Loe. Yorkshire. Arn. ceras. Fig. 20 and 20a, specimen with incomplete living chamber, a very broad abdomen, and deep channels, loc. Semur. Arn. tardecrescens. Fig. 21 and 22, sutures of a specimen of a normal form (abdomen narrower than the last), at the diameters of 63 mm. and 83 mm. Loe. Semur. Arn. Bodleyi. Fig. 23, broken specimen, showing planorbis-like folds on the young shell, with living chamber incomplete. Fig. 24, more involute variety with similar fold in the young and incomplete living chamber. Fig. 24a, suture of the same, enlarged. Loc. Semur. Arn. falearies. Fig. 25, young with slight tubercles on the genicule. The last quarter is part of the living chamber. Fig. 26, larger specimen, ribs not tuberculated or arising from, tubercles as in the above, and keel very prominent. Fig. 27, cast with deep channels and abdo- men very narrow. Loc. Semur. Arn. kridioides. Fig. 28, specimen showing the smooth young, the early period at which the pile. begin, and the similarity of the umbilicus to that of a normal species of Arnioceras. Loe. Basle. 1 See Plate I. Fig. 15, 16. 2 See Pl. Til. Fig. 1, la. GENESIS OF THE ARIETIDAE. PLATE Il. )DW ROETTER & HYATT, DEL. HELIOGRAPH, HART & VON ARK, N. ¥ ee iy rb Alga PLATE III. Arn, Hartmanni.' Fig. 1, 1a, cast. Fig. 1 shows the thick shell lying on the keel of the cast. Loc. Lyme Regis. ; Cor. kridion. Fig.2, young. Fig. 2a, outline of young suture of Fig. 2, enlarged. Fig. 3, east. The last volution in this represents the living chamber. Loc. Balingen. Cor. rotiforme, var. A. Fig. 4-4a, young smooth stage and suture. Tig. 5, 6, 8, 8a—10, 10a, older stages of same. Fig. 7, 11-13, pathological cases with pile crossing the abdomen. Loc. Semur. Fig. 14, 15, 15a, 16, kridion-like variety of this species. (Hig. 14, loc. Stuttgardt ; Fig. 15, 16, loc. Balingen.) Fig. 17, 17 a, variety with discoidal form and large tubercles, figure reduced to about two thirds, loc. Semur. Fig. 17 b, sutures. Cor. Bucklandi, var. sinemuriense. Fig. 18, figure reduced to less than one half, show- ing the divided pile of the young (sinemuriense stage), and the solid Bucklandian pile of the adult, loc. Semur. j Cor. latum. Fig. 19, young with narrow abdomen. Fig. 22, older stages of same variety, showing affinities for Cor. rotiforme and Bucklandi. Fig. 20, section of young of variety with broader abdomen. Fig. 21, 23, 23a, older stages and suture of same. Loc. Semut.? ! See Plate II. Fig. 17. 2 These specimens all appear to be in the nealogic stages of development. PLATE Ill. GENESIS OF THE ARIETIDAE. HELIOGRAPH, HART & VON ARX, ROETTER & HYATT, DEL. i PLATE IV. Cor. lyra.t Fig. 1, young of variety with broad abdomen and closely arranged pile. Fig. 2, young of var. B, with narrower abdomen and rounded sides. Fig. 3, older stage of same variety. Fig. 4, 5, nearly full-grown stage of same variety. Loc. Semur. Fig. 6, 7, full-grown stage of variety with narrower abdomen, flattened sides, and closely arranged pile, figure reduced to about one half, loc. Filder. Fig. 8, young of same variety as Fig. 1, but transitional to var. B. Fig. 9-11, young of var. C. Fig. 11a, suture of same. Fig. 12-14, young of variety like Fig. 1, 1 a, and Fig. 8, but with narrower abdomen and more flattened sides. Loc. Semur. Fig. 15, 16, old specimen of var. C, with very convergent sides, and tubercles considerably ane diameter 250 mm., loc. Gmiind. Fig. 17, sutures on the inner side (dorsum),? loc. emur. 1 See Plate V. Fig. 1-3. 2 All the specimens figured were casts. ee ee . GENESIS OF THE ARIETIDAE. ROETTER & HYATT, DEL. PLATE IV. HELIOGRAPH, HART & VON ARX, N. Y. PLATE V. Cor. lyra.1 Fig. 1-3, half-grown shell of same variety as Fig. 6, 7, Plate IV., showing how narrow the abdomen is in some specimens before the shell was full grown. Fig. 5a, suture, slightly older than that delineated on Fig. 2, showing the changes which had taken place. Loe. Semur. Cor. Gmuendense.? Fig. 4, half-grown specimen, exhibiting variety with discoidal form and compressed whorls, figure reduced to less than one half. Fig. 5, full-grown specimen of same variety with marks of approaching old age upon the last whorl, figure reduced to less than one half. Loc. Semur. Fig. 6, specimen in which the wider abdomen and young proportion of Fig. 4 were maintained until a much later stage than in Fig. 4 or 5, figure reduced to about one half. Fig. 7, umbilicus of Fig. 6, about natural size, to show the smoothness of the young whorls. Loc. Aargau. Fig. 8, 9, same variety as Fig. 5, but showing the effects of senile decline in the convergence of the sides, degeneration of the pile and tubercles, diameter 205 mm., loc. Semur. 1 See Plate IV. 2 See Plate VI. GENESIS OF THE ARIETIDAE. PLATE V. ROETTER & HYATT, DEL. HELIOGRAPH, HART & VON ARX, N. Y. el Oa iY PLATE VI. Cor. Gmuendense. Fig. 1, 2, figures reduced to about one third, taken from a specimen which exhibited compressed whorls and convergent sides at an older stage than in the specimens figured on Plate V. Old age is apparent also in the two closely approximated sutures of outer whorl. Loc. Aalen. Cor. trigonatum.! Fig. 3, large variety, having stouter and more numerous whorls, and arriving at the same stage of senile decline later than in the smaller variety described. in the text, figure reduced to about one third. Loc. Aalen. Cor. Sauzeanum.? Fig. 4, young, loc. Whitby. Fig. 5, another older specimen, loc. Semur. Fig. 6, another specimen, broken across, showing embryo, and having keel just begin- ning to be perceptible on the outer whorl. Fig. 7, centre of same section enlarged, showing young whorls. Fig. 8, specimen like Fig. 12,in having no keel on the outer whorl. Fig. 9, variety in which both tubercles and keel appear at an earlier stage. Loc. Whitby. Fig. 10, 11, same variety, showing stouter form of young and earlier developed keel. Fig. 12, 13, D’Or- bigny’s type, showing the prolonged duration of the smooth stage, and absence of the keel. Fig. 14, adult of var. Gaudryi. Loe. Semur. Cal. raricostatum.? Fig. 15, Johnstoni-like variety, loc. Balingen. 1 See also Plate VII. Fig. 1, for side view. 2 See Plate VIII. Fig. 1-3. 3 See Plate I. Fig. 24, 25. PLATE VI. GENESIS OF THE ARIETIDAE. HELIOGRAPH. HART & VON ARA&, N. Y. ROETTER & HYATT, DEL. PLATE VII. Cor. trigonatum. Fig. 1, old specimen of the stout variety, showing adult suture and the closer approximation and degenerative changes in lobes and saddles in old age, figure reduced to about one third. Loc. Aalen.? Cor. bisuleatum. Fig. 2-4, young. Fig. 5, 6, young with earlier developed and more prominent pile. ‘The coarse heavy pile of the last figure are remarkable, and the young has also a broader abdomen and a form like an older stage of Cor. latum. Fig. 7 shows the decrease in breadth of the abdomen with age, which is also seen in Fig. 6. Fig. 8, older stage, with suture. Fig. 9, 10, nearly full-grown stage of same variety. Loc. Semur. Agas. Scipionianum.? Fig. 11, 12, young of gibbous variety. Fig. 13, 14, young, show- ing development of pile from folds. Fig. 13a, 14a, enlarged sutures also indicated upon corresponding figures. Fig. 15, front view of same specimen as that of Plate X. Fig. 13. Loe. Semur. 1 See, for ventral view of same, Plate VI. Fig. 3. 2 See Plate X. Fig. 11-13. Summ. Pl. XIII. Fig. 7. GENESIS OF THE ARIETIDAE. ROETTER & HYATT, DEL. PLATE VII. HELIOGRAPH, HART & VON ARX, N. Y. ata east ae se) PLATE VIII. Cor. Sauzeanum.! Fig. 1-3, var. Gaudryi, older and younger whorls of the same speci- men, loc. Leicestershire. Fig. 3a, dorsum showing sutures. Aster. obtusum.? Fig. 4, 5, young of var. EH, showing large tubercles and broad abdo- men. Fig. 6-8, older stage of same specimen. loc. Lyme Regis. Agas. levigatum. Fig. 9, side view of var. B, with living chamber. Fig. 10, var. D, sec- tion showing the extremely broad young and secondary helmet-shape of the later whorls, which are like those of Psiloceras except of course in the keel (the outline of the centre is uncer- tain except as regards the breadth). Fig. 11, var. A, showing the absence ofa keel. Fig. 12, var, D, showing outline of aperture and living chamber, loc. Semur. Fig. 13, very stout young, showing the goniatitic form, and striations like those of Agas. striaries. Fig. 14, section of variety from Lyme Regis.® 1 See Plate VI. Fig. 4-14. 2 See Plate IX. Fig. 1. 3 All these figures are enlarged about two diameters, except Fig. 9 and 12, which are about natural size. GENESIS OF THE ARIETIDAE. PLATE VIII. ROETTER & HYATT, DEL. HEAIOGRAFH. HART & VON ARA, N. Y- ran eS mes aie aes PLATE IX. Aster. obtusum.'’ Fig. 1, var. D, loc. Lyme Regis. Aster. stellare. Fig. 2, 3, figures reduced to about two thirds, loc. Tiibingen. Aster. acceleratum.’ Fig. 4 shows variety having the nearest approximation to Aster. stellare. The section is similar to that of Fig. 9, but is more involute. Loc. Semur. Aster. Brooki. Fig. 5, 6, stout-whorled, broad-abdomened variety, approximating to Turneri, but with broader sides at the same age. Fig. 7, older stage of the same variety. Figures reduced to about two thirds. Loe. Lyme Regis. Aster. Turneri. Fig. 8, 9, old specimen, figures reduced to about two thirds, loc. Semur. Aster. Collenoti.? Fig. 10, young, showing compressed form and acute abdomen. Fig. 10a, b, the same more enlarged, but the figures fall short in depicting the acuteness of the abdomen, and by improper shading show a keel which has no existence. Fig. 11, young and older stage; the young are again too rounded, the outer whorl is however approximately accu- rate. Fig. 11a, b, centre of same more enlarged, showing the involute form of even this early stage, but the front view is not sufficiently compressed. Loc. Semur. Agas. Scipionis. Fig. 12, 13, lateral and sectional view of young, loc. Semur. Agas. striaries. Fig, 14, 15, variety with flattened sides and broad abdomen, loc. Semutr. 1 See Plate VIII. Fig. 4, 8. 2 The umbilical shoulders are not made abrupt enough in this figure, and the umbilicus is shallow as in Aster. stellare, instead of being deep as in this species. See Plate X. Fig. 3. 8 See Plate X. Fig. 10. = re » = Z = F ime 4 2 [o} g a bk c S = ae x zt c oO 9 g Wl =r ui —¢ a . = Lu (ae —t a . SE i E 4 ° > oe B cy ny c Ww fe im F Ww fe) © @ SUMMARY PLATE XIII. SUMMARY PLATE XIV. The three preceding plates do not illustrate the biological relations of the Aietidee as a whole with sufficient clearness, and this plate has been added for the purpose of supplying the defi- ciency. ‘The series of Psiloceras has been placed in what may be deemed its true position, be- tween the Plicatus stock and the Levis stock; otherwise, the arrangement is the same. ‘The resemblances of the morphological equivalents in each series can be readily seen by following the forms along horizontal lines from left to right. The independence of the origin of these representative forms can be studied by following up the series in vertical lines, which represent descent. To a large extent, also, the more obvious differential characters which distinguish each series become appreciable by the same process. Psil. planorbe, var. leve, Fig. 1; var. plicata, Fig. 2. Schlot. catenata, Fig. 3, is the radical of this series. Schlot. angulata, Fig. 24, is evidently a transition to the next species. The artist has exchanged Fig. 4 with Fig. 24. Schlot. Charmassei, Fig. 5. ‘The whorl is more involute, but the degenerate characters of compression in the whorls and shallowing of the abdominal channel begin to appear. Schlot. Boucaultiana, Fig. 6. The involution has attained its maximum, and the degen- eration of the pilz and channel is well marked. Weh. curviornatum (sp. Wih.), Fig. 7, is undoubtedly distinct from Schlot. angulata, and is one of the radicals of this series. Weh. haploptychum (sp. Wih.), Fig. 28. The artist has exchanged Fig. § with Fig. 28. Weh. toxophorum (sp. Wih.), Fig. 9, is a degenerate shell, having compressed whorls, and pilee crossing the abdomen, as in the proximate radical Weh. curviornatum. It is, however, more involute. Weh. Emmerichi (sp. Wah.), Fig. 10, shows a notably involute shell, with degenerate pilz and compressed whorls. Cal. tortile, Fig. 11, is the radical of this series. Cal. carusense, Vig. 12, has similar young to that of tortile below. Cal. Nodotianum, Fig. 13, is very similar to carusense, but with more compressed whorls and better developed pile. ‘ Cal. eyeloides (sp. Wih.), Fig. 14, shows compressed degenerate whorls. Cal. Castagnolai (sp. Wah.), Fig. 15, is more degenerate than the last, but slightly more involute. Cal. abnormilobatum (sp. Wih.), Fig. 16, is a dwarfish and more degenerate form than Castagnolai, but has more involute whorls. Cal. laqueum, Fig. 17, is an extreme form of this species, which approximates very closely to a true spiratissimum. This figure is therefore placed to the right, and under Verm. spiratissimum. Verm. spiratissimum, Fig. 18, shows typical form, with but slight channels. Verm. Conybeari, Fig. 19, shows normal untuberculated variety, with stout whorls and deep channels. Verm. ophioides, Fig. 20, exhibits the tuberculated pile of this species. Psil. aphanoptychum (sp. Wih.), Fig. 21, is one of the Plicatus stock of Psiloceras. Psil. Kammerkarense (sp. Wah.), Fig. 22, shows the more involute and plicated form of this subseries. Psil. mesogenos (sp. Wah.), Fig. 23, is an involute shell belonging to the true Levis stock. Arn. semicostatum, Fig.4. The figure represents the nearly full-grown shell; but if the keel were absent, the smooth whorls of the young would closely resemble the adult whorls of Psil. planorbe, var. leve. The artist has exchanged Fig. 4 with Fig. 24. Arn. Hartmanni, Fig. 25, exhibits young and adult characters like those of the preceding. Arn. tardecrescens, Fig. 26, belongs to another subseries of forms than that in which it is placed, but it serves to show that quadragonal whorled shells with channelled abdomens existed in this genus. 1 1 Two subseries ought to have been shown here, but in trying to reduce the size of the plate the forms have been placed in the same line. A similar liberty has been taken with the subseries of Caloceras and Arnioceras, but this does not interfere with the truthful presentation of the general zodlogical relations of the forms. SUMMARY PLATE XIV. (continued). Arn. Bodleyi, Fig. 27, shows a slightly degenerate compressed whorl, and is the terminal form of the subseries containing Hartmanni. Arn. kridioides. Fig. 28 gives a view of the transition between Arnioceras and the lowest species of Coroniceras. The smooth young straight pilz and divergent sides of the adult whorl are clearly shown. The artist has exchanged Fig. 8 with Fig. 28. Cor. Sauzeanum. Fig. 29 shows the later nealogic and ephebolic stages, having the peculiar divergent sides, flattened abdomen, and prominent tubercles of a typical coroniceran form. The young, however, still retain the smooth aspect, indicating derivation from Arnioceras. Cor. rotiforme. Fig. 30 represents a form similar to Cor, coronaries. Cor. Lyra, Fig. 31. This is as a rule much smaller than rotiforme. The sides are more convergent, and the whorls more compressed and less numerous than in that species. Cor. trigonatum, Fig. 32, exhibits the effects of the premature development of old age characters. Fig. 1 on the extreme right shows the dwarfed form of Psil. planorbe, var. leve, from which both the arnioceran as well as the agassiceran series may have been derived in Central Europe. Agas. levigatum. Fig. 33 shows the more compressed variety of this species. Agas. striaries, Fig. 34. The striations were too fine to be represented. Ast. obtusum. Fig. 2 shows the stouter variety with well marked channels with stout gibbous whorls and broad abdomen. This has young almost identical with the adults of the stout varieties of Agas. levigatum. Ast. Turneri. Fig. 36 shows typical variety, with flattened sides and deep channels, Itis notably more involute than obtuswm. Ast. Brooki. Fig. 37 shows an extreme involute variety of this species, with very conver- gent sides and narrow abdomen. ‘The channels are almost obliterated, and the keel very prominent. Ast. Collenoti. Fig. 38 gives a view of this remarkable dwarfed form, in which degenera- tion of the pile and the channels and convergence of the sides have produced morphological equivalence with Oxyn. oxynotum and Guibali. The amount of the involution is greater than in any preceding species of the same series. Agas. Scipionianum. Fig. 39 shows the stouter, heavily tuberculated variety, which has young almost identical with the stouter varieties of Agas. striaries. Agas. Scipionis. Fig. 40 shows an aged specimen in the Museum of Comparative Zoology, with extremely involute whorls, but keel still prominent. The degeneration of the adult as regards the pile and form can, however, be inferred from this figure. The old of Scipionianum at the same age is much less changed, and does not exhibit increased involution of the whorls. Oxyn. oxynotum, Fig. 41, 42. The first figure shows the young of a variety in which at an early stage there is close likeness to the young of Agas. striaries, and the adults of Agas. levigatum. Oxyn. Simpsoni. Fig. 43 shows the stouter form and slightly greater involution of the whorls in this species when compared with oxynotwm. Oxyn. Lymense. Fig. 44 shows the greater involution of whorls as compared with any preceding form of the same subseries, and the very acute degenerate whorl. Oxyn. Greenoughi. Fig. 45 shows the stout form of the whorls better defined, and pile of this subseries as compared with the oxynotum subseries. Oxyn. Lotharingum. Fig. 46 shows the smaller size of this species, and the degeneration of the pile. The involution of the whorls is, however, greater than in any preceding species.? Oxyn. Oppeli. Fig. 47 shows the extremely involute form of the Middle Lias. The stout whorls indicate that no great amount of degeneration had taken place. It may have been a direct descendant of Gieenoughi. 1 The extreme old age of this form is marked by decrease in the amount of involution of the whorl, and also by the loss of the prominent hollow keel. RIX EB2LWId AMYWANGS NOLSOg ‘SH3INIYd P SYSAVYONA “OD xN7 a ' 1 ¥ . s ‘ Ar _ a PS iy ‘ en ‘ yar. ¢ al ae ae ste _ vn : 3 9088 014