Issued October 10. 1910. . S. DEPARTMENT OF AGRICULTURE, BUREAU OF ANIMAL INDUSTRY. BULLETIN 125, PART 1. A. IX MELVIN, CHIEF OP BUREAU. GID PARASITE AND ALLIED SPECIES 9 ~~~ OF THE CESTODE GENUS MULTICEPS. I. HISTORICAL REVIEW. BY MAURICE C. HALL, Junior Zoologist, Zoological Diri L J E r : WASHINGTON : GOVERNMENT PRINTING OFFICE. 1910. - N X THE BUREAU OF ANIMAL INDUSTRY. r.hi.f: A. D. MEL YIN. Assistant Chief: A. M. FARRINGTON. Cfiiif CJcrl-: CHARLES ('. CARROLL. al Husbandry Division: GEORGE M. ROMMEL, chief. Biocht.;iir ftirision: M. DORSET, chief. Dairy D^qion: B. II . RAWL, chief. Inspection Division: RICE P. STEDDOM, chief; MORRIS WOODEN, R. A. RAMSAY, and ALBERT E. I' I; -ITXKE, associate chiefs. Pathological Division: JOHN R. MOHLER, chief. Quarcmli/tc ]>irisiori* RICHARD W. Jin KMAN, chief. Zoological Division: B. H. RANSOM, chief. iment Station: E. ( . SCHROEDER, superintendent. Editor: JAMES M. PICKI /.OOLOCICAI, DIVISION. Cl/uf: I!. II. HANSOM. AKX/xtflnl Zonlnfj'ixt: Al.HKRT .1 1.VSSA I.I.. Junior Zoologists. HAHIIY \\'. (IHAYHII,!., M.-vruici-; ('. HALL, HOWARD <'KA\VI.I:Y, and WINTHROP D. FOSTER. Issued October 10, 1910. U. S. DEPARTMENT OF AGRICULTURE, BUREAU OF ANIMAL INDUSTRY. BULLETIN 125, PART 1. A. D. MELVIN, CHIEF OF BUREAU. THE GID PARASITE AND ALLIED SPECIES OF THE CESTODE GENUS MULTICEPS. I. HISTORICAL REVIEW. BY MAURICE C. HALL, Junior Zoologist, Zoological Division. WASHINGTON: GOVERNMENT PRINTING OFFICE. 1910. LETTER OF TRANSMITTAL. U. S. DEPARTMENT OF AGRICULTURE, BUREAU OF ANIMAL INDUSTRY, Washington, D. C., June 16, 1910. SIR: I have the honor to transmit herewith, and to recommend for publication as a bulletin, the accompanying manuscript entitled "The Gid Parasite and Allied Species of the Cestode Genus Multiceps. Part 1. Historical review," by Maurice C. Hall, of the Zoological Division of this Bureau. Mr. Hall has been making a most comprehensive study of gid, and his investigations will furnish an important contribution to our knowl- edge of this deadly disease of sheep, which has only in recent years been recognized as established in the United States, the first definite evidence of its presence as an enzootic having been published in 1905 in Bureau of Animal Industry Bulletin 66. It is intended to publish later, as succeeding parts of the present bulletin, the results of Mr. Hall's investigations, now in progress, con- cerning the morphology and life histories of the parasites in question, as well as the symptomatology, treatment, prophylaxis, etc., of gid. Respectfully, A. D. MELVIN, Chief of Bureau. Hon. JAMES WILSON, Secretary of Agriculture. CONTENTS. I'age. Introduction 5 Multiceps multiceps 6 Historical sketch 6 Gid in the United States 16 Gid in Canada 29 The hosts and occurrences of the larval Multiceps multiceps 30 The occurrences of the adult Multiceps multiceps 41 Economic importance of gid 42 Alleged causes of gid 46 Names applied to gid and giddy animals 47 Common names of the gid parasite 49 Synonymy 50 Multiceps serialis 56 Historical sketch 56 The hosts and occurrences of the larval Multiceps serialis 58 The occurrences of the adult Multiceps serialis 03 Economic importance 64 Synonymy 65 Multiceps lemuris 66 Historical sketch 66 Synonymy 66 Multiceps polytubcrculosus 67 Historical sketch 67 Synonymy 67 Multiceps spalacis 67 Historical sketch 67 Synonymy 67 Cysticercus botryoidcs 68 Historical sketch 68 Synonymy 68 Acephalocystis ovis tragclaphi 68 Historical sketch 68 Synonymy 68 ILLUSTRATION. FIG. 1. Map of Montana, showing distribution of gid in sheep. THE GID PARASITE AND ALLIED SPECIES OF THE CESTODE GENUS MULTICEPS. PART I. HISTORICAL REVIEW. INTRODUCTION. Coenurus is the name commonly applied to a larval cestode group of considerable importance to helminthologists from a historical and scientific standpoint, for it was with one of its species, commonly referred to as Cc&nurus cerebralis, that Steenstrup's theory of the alternation of generations was first completely demonstrated for cestodes by Kuchenmeister, who, in 1853, produced the adult cestode or tapeworm in the primary host by feeding the larval form to the dog, and produced the larval cestode or bladderworm in the secondary host by feeding the eggs of the adult tapeworm to the sheep. This work of Kuchenmeister's and that of Von Siebold along the same line is taken by Braun (1894a), a in his classic work on cestodes, as marking the beginning of the fourth and latest period in helminthology, dating from 1851. This same species, C. cerebralis, is of considerable economic interest to veterinarians and stock raisers, and especially to sheepmen, as being the cause of the disease commonly known among English- speaking people as gid. In spite of the fact that the disease caused by this parasite, as well as something of its nature, was probably known in the fourth and fifth centuries B. C., and that the parasite itself was observed at least as early as 1634 A. D., its parasitic nature known since 1780, and its life history known for over half a century, there are still some mistaken popular ideas about it, and also some errors, disagreements, and uncer- tainties in the writings of scientists as to the specific identity of this and various other forms of coanurus that have been described from different hosts, and also as to the correctness with which certain par- a Bibliographic citations refer, wherever possible, to Stiles and Hassall's (1902-19 ) Index-Catalogue of Medical and Veterinary Zoology, Authors, Bureau of Animal Indus- try Bulletin 39, United States Department of Agriculture. References not in Bul- letin 39 are indicated by the use of Greek letters and will be covered in a supplemental bibliography,, to be published later. 5 6 THE GID PARASITE AND ALLIED SPECIES. asites are listed from certain hosts. The writer has endeavored to correct some of these errors in this paper, and it is proposed in a series of papers to give a comprehensive account of the cestodes having a coenurus larva. The first form to be considered is the brain bladderworm of sheep, usually known as Cwnurus cerebralis, but which, as will be shown, should be known by the name Multiceps multiceps, proposed here for the first time. In this article the word " cosnurus " will not usually be capitalized ; it will be used merely as the name of a larval stage, like the words " cysticercus," "cercaria," "leptocephalus," etc. It is not entitled to be used as a generic or subgeneric name, owing to the pri- ority of Multiceps, but as it is still much more commonly used in this way than Multiceps, and as reference must be constantly made to quotations where it is used in combination with some specific name, especially in the form Ccenurus cerebralis, it will often be clearer to use this form instead of the correct one. MULTICEPS MULTICEPS. HISTORICAL SKETCH. Braun (1894a) makes his first period in helminthology cover the work of antiquity and the middle ages up to 1600, and in the litera- ture of this period, relatively barren from a scientific standpoint, almost no references are to be found that can be construed as refer- ring to gid. However, a disease like gid, involving, as it does, a deli- cate arrangement of alternating hosts, must have existed long before primitive man passed from the hunting to the pastoral stage. It is not the sort of disease to arise by rapid facultative adjustment or out-of-hand adaptation. The very fact that gid exists to-day is proof enough in a disease of this sort that it existed thousands of years ago. Undoubtedly, in the days when the ancestral dog pur- sued the wild sheep, the nice adaptation of a brain parasite that would interfere with muscular activity and blunt the sense perceptions, making flight and escape difficult, must have furnished a striking example of a life habit well calculated to perpetuate a parasite, but it could scarcely have been more satisfactory than the new arrange- ment introduced by man when he domesticated the sheep and put its former enemy, the dog, in charge of it to run over its pastures as a constant companion and to eat the discarded heads and diseased brains of giddy sheep an enemy still. A prolonged search of ancient literature would no doubt show some references which might readily be taken as descriptions of gid. The symptoms are so striking that pastoral peoples, like the Arabs, Jews, and Greeks, must have noted and described them ; but finding such references involves a tedious search and more lime than ran profitably be spent on the work. HISTORICAL SKETCH OF MULTICEPS MULTICEPS. 7 One such reference occurs in Kuhn's edition of Hippocrates (1825), who is believed to have lived 460 to 375 B. C. The follow- ing is quoted from Adams's translation of Hippocrates (1886or), describing excess of fluids on the brain in epilepsy: This you may ascertain in particular, from beasts of the flock [i. e. , sheep] which are seized with this disease, and more especially goats, for they are most frequently attacked with it. If you will cut open the head you will find the brain humid, full of sweat, and having a bad smell. It is, of course, impossible to make a positive statement of fact on anything less than complete and accurate observations. Obvioush* there was no one in the tune of Hippocrates who could be expected to make and record such observations in a case of gid, and existing editions of Hippocrates are open to the suspicion of having in them observations not properly referable to Hippocrates. Hence we can not say certainly that Hippocrates actually saw cases of gid, but on the strength of the reference given, agreeing as it does with the certainty that gid among sheep must have existed for ages, it is fair to state that Hippocrates probably saw cases of gid four or five centuries before the Christian era. The fact that the brain of sheep was found full of fluid points, among other things, to hydrocephaly, which may follow the invasion of the gid parasite, according to Miiller (1877a), or to the gid parasite itself. Gid probably was not rare in those days when sheep were everywhere tended by dogs and the prophylaxis of the disease was undreamed of. The "bad smell" may have been due to delay in post-mortem examination, to hydro- cephalus purulentus as a sequel of gid, or it may easily have been noted hi the ccenurus vesicle, as my own observations show that the coenurus fluid serves as an excellent medium for decomposition bacteria, the odor of the fluid in a graduate becoming intolerable in twenty-four hours at ordinary room temperature. Guetebruch (1766a), according to Kuchenmeister (1880a), states in an article on gid that when perforation of the skull occurs, as it sometimes does in gid, the brain decomposes and becomes purulent, the brain and bone marrow turning to water and becoming putrid. The writer has never seen such a case, but it is evident that if the perforation of the skull were followed by perforation of the skin as well, it would afford entrance to bacteria, with possibly a result similar to the one given. Finally, the fact that these post-mortem findings are given for sheep suffering from "the sacred disease," a term covering epilepsy and other brain disorders, would indicate the possibility of gid, as the symptoms of nervous disturbances are very marked in this disease. Adams, the translator of Hippocrates from whom the foregoing quotation is taken, and himself a physician, refers to the lines quoted as follows: It is well known that this is also the case with sheep, and that they are subject to the disease called the sturdy [i. e., gid], which is indisputably a sort of epilepsy. 8 THE GID PARASITE AND ALLIED SPECIES. In the somewhat limited literature on helminthology for the period from 1600 to 1800, Braun's (1894a) second period, the gid parasite figures to a proportionally large and increasing extent. The citations from this period are given rather fully, as they are in works which are not readily available to many. In the first part of the nineteenth century, Braun's (1894a) third period, there are numerous references to gid, and since 1850 and the work of Kuchenmeister, which was done soon after, not a year has passed in which few to many notes on the brain bladderworm, its adult tapeworm, or its effects, have not appeared. This increase in the amount of literature is perhaps concomitant with an increase in number and distribution of sheep and cases of gid, as well as with increasing knowledge of the parasite. In general the large amount of literature is due to the attractive combination of scientific and eco- nomic interest which has induced many persons to publish notes on the disease and its parasite from one or both standpoints. The early notes on cosnurus deal only with Ccenurus cerebralis (= Multiceps multiceps) and especially with the disease caused by it. It was nearly two hundred years after Scultetus (1672a) had seen the first unmistakable case of gid that I have found recorded, before the first ccenurus which we may regard as other than C. cerebralis was noted by De Blainville (1828a). Scultetus saw his case in 1634. The first available note published during Braun's (1894a) second period of helminthology dealing with C. cerebralis is that of Rolfinck (1656a) who, in a work on medical anatomy, writes of vesicles full of water and humor in the third ventricle of sheep as the cause of a vertigo. This may be safely accepted as a reference to C. cerebralis. The description is in general terms just the one a casual observer would give of this parasite, as witness the statement of a correspondent to the veterinary editor of a periodical (Vet. Ed. Amer. Shepherd's Bulletin 1903?-) to the effect that he found hi a sheep's head "a bag of water which burst and ran out when I pressed upon it." The next available article on the subject of gid published during this period is that of Wepfer (1658a). The part relating to C. cerebralis gives at this early date notes on the characteristic symptoms of the disease, its pathology, and the morphology of the water bladder. The disease is further recorded as a frequent cause of death in cattle, and the peasants are credited with a form of operation involving percussion and surprisingly good for that date. Heusinger (1853a) quotes from a work of Bartholinus (1667aO, not available to me, a statement of a species of frenzy and vertigo which in 1661 attacked horses, cattle, and sheep, and notes that worms were found in the heads of the animals attacked. These cases may have included, and very likely did include, cases of gid. The next available article dealing with C. cerebralis is that of Scultetus (1672a), who in a Latin treatise on surgery gives the HISTORICAL SKETCH OF MULTICEPS MULTICEPS. 9 description of a case seen at the earliest date at which we have found a case recorded. The following is quoted from an English translation of the same work (Scultetus, 1674a): Observation X. Of a Vertigo in a Sheep, proceeding from an Abscess in the Brain. In the Year 1634, December the 24th. Being in the shop of Nicolas Kite he made mention of his sheep, among which one was troubled with a Vertigo, or Giddiness, the Germans call it Wirbling: this Disease one who dealt in sheep affirmed to be inci- dent to the fairest of the Flock; that hereby their whole Brain would be turned into Water and then they would fall down dead on a sudden. The Chyrurgion therefore commanded that one of those sheep which was weakened with this Giddiness, and turning around, should be killed, and sent me the head. Scultetus found nothing in the ventricle. Afterward I lifted up the organs of smelling * * * and on the left -side, between the Brain and the Pia mater, I found an abscess, like the Bladder of a Fish, full of very clear water * * * I wondered that * * * the sheep should not labour under an Apoplexy, or a Palsy, rather than a Vertigo. / In 1645 Scultetus lost a sheep by the same disease, and in the work just noted writes: I dissected the Head * * * and presently on the left-side as it were of the backward part of the Head, under the Dura Mater, I found a Bag of the thickness of a Fisches Blader, filled with Water, and little Worms, such as are bred in Cheese; for it began to putrefie at the bottom. This Coated Tumour being bigger than a Hens Egg, had so insinuated itself into the substance of the Brain, that it did somewhat press upon the third Ventricle. This Sheep, as the Shepheard reports, turned herself round about towards the night & all that day she dyed. That gid was not uncommon in the seventeenth century is clear from the fact that Rolfmck (1656a), writing of vertigo, refers to it as occasionally (nonnumquam) caused by sacs of water on the brain in sheep. Wepfer (1658 a) notes it as a serious and common disease of cattle in Switzerland. In the account of Scultetus (1674a) it appears that a sheep dealer recognizes the disease as one common enough in Germany at that time to have a colloquial name, "Wirbling." The next reference to gid is by Wepfer (1681 a) and is identical with the one already given, being in a later edition of the original work of 1658. Kuchenmeister (1880a) refers to an article by Brunner (1694or), not available to me, and quotes from it a statement to the effect that Brunner had dissected the head of a giddy calf, "vituli vertiginosi," and in the cerebral substance had found three hydatids the size of pigeon eggs and full of limpid fluid. Kuchenmeister takes this to refer to Casnurus cerebralis, which it obviously does. a The original Latin text reads "in tonstrino Nicolai Reutte." The translator has translated not only the text but also the proper names, rendering the German name Reutte by its English equivalent, Kite. 6 This last statement should read " towards the right," the Latin word here being ''dextram." 51674 Bull. 125, pt. 110 2 10 THE GID PARASITE AND ALLIED SPECIES. The next reference is in Wepfer (1724a). The first two parts of this article consist of the two parts making up the edition of 1658. With these is incorporated a third part. The same references to gid occur in the parts already published and referred to above. In the new part is a new reference to hydatids in the brain of cattle as being commonly believed to be the cause of the vertigo accompanying them. He has seen the peasants perforate the skull and extract these in operations and has also seen the hydatids demonstrated post-mortem. Hoffberg (1759a), in a dissertation on Cervus tarandus, first presented in 1754, writes under the heading of diseases of this animal, of a vertigo or "Ringsjuka" causing the reindeer to turn in circles. Braun (1894a) takes this as a reference to C&nurus cere- bralis, which is a perfectly reasonable assumption. The presence of the parasite in the reindeer, however, is unsupported by post- mortem evidence in this reference, and, so far as I am aware, such evidence is lacking in any subsequent writings. The occurrence of the gid parasite in the reindeer must therefore be considered doubt- ful. It seems the more doubtful hi that Brehm (1877^) states that reindeer are attacked by the larva of a gadfly, specified by Moniez (1880a) as Cephenomya trompe, which penetrates from the nasal cavity to the brain, causing a fatal "Drehkrankheit" or gid, and it may have been this disease, apparently a common one, which Hoffberg saw. Kuchenmeister (1880a) quotes from a treatise on diseases of sheep by Guetebruck (1766^), already noted as not available. In this treatise it is stated that the disease attacks lambs and yearlings, but not old sheep; that some are born with it; that a water bladder forms on the brain and may penetrate the skull; that when the disease has not gone too far the flesh may be used and the head and feet thrown away [very bad advice], but if the disease has gone too far the entire carcass should be done away with. As a method of treatment he gives venesection on the temple and nose. Stier (1776a) has an article on gid, of which only the review was seen by me, the original (Stier, 1775a) not being available. The article takes up a long list of supposed causes of gid and rejects them, the water bladder in the head being held guilty of causing the trouble. Stier also draws a careful distinction between actual gid due to C. cerebralis and simulated gid due to the presence of (Estrus larvae in the nostrils, the latter presenting the symptoms most com- monly mistaken for gid. According to footnotes in Bloch (1780a), Hastfer (I776a) and Ranstler (1776<*) have published references to gid, but these are not available. Bloch states that they attributed gid to the bladder on HISTORICAL SKETCH OF MULTICEPS MULTICEPS. 11 the brain, and that Ranstler was the first to notice the small bodies on the bladder and surmised that worms arose from them. According to Braun (1894a) and others, the cestode nature of the water bladder found in the brain of giddy sheep was first pointed out by Leske (1780a) and by Goeze (1780a), independently. These references are not available to me. Braun notes that Goeze recog- nized the cestode heads and considered them as the embryos of the bladderworms which are found in the omentum and liver of sheep and swine. He also notes that Leske found Tsenia multiceps (=Coenurus cerebralis), recognizing the characteristic hooks and suckers. Kuchenmeister (1880a) quotes part of Leske's article showing that Leske made a very careful study of the morphology and pathology of the parasite. He noted the heads invaginate and evaginate through the bladder wall. From the presence of so many of these heads, he observes that we may consider the animal as many tapeworms attached to a common bladder, or as one tape- worm with many heads. Hence it would be appropriate to call it the many-headed tapeworm, so he names it Tsenia multiceps. This last is important, as it establishes the fact that the correct specific name of the gid parasite is multiceps. The preceding note from Braun (1894a) confirms the correctness of Kiichenmeister's (1880a) quotation, and in addition Mr. Sherborn has very kindly verified the reference in the library of the British Museum. It appears from evidence to be considered later that Leske's work antedates that of Goeze in the same year. Were it otherwise, Goeze's article need not be considered, as, according to Braun's synopsis, he regarded the heads of the parasite as the embryos of the bladder- worms found in the omentum of sheep and swine, and hence pre- sumably proposed no new name for the brain parasite, as there would be no reason for it under the circumstances or a proper appli- cation for the name had he done so. In a discussion of the synonymy of this parasite, Stiles and Steven- son (1905a) accept as the specific name the one proposed by Bloch (1780a). Bloch makes the genus Vermis vesicularis for the bladder- worms, and divides these into three species, of which Vermis vesicu- laris socialis is the brain bladderworm of sheep. But though this article of Bloch's bears the same date as those of Leske and Goeze, viz, 1780, Leske's article is nevertheless older, and the name pro- posed by him is therefore entitled to priority. This is evident from Bloch's own article, which shows that Bloch had read Leske's article of the same year. Bloch states that Ranstler first noticed the small bodies on the bladder walls and surmised that worms arose from them, but that Leske and Goeze observed that these bodies were actually bladderworms. He states that Leske has described them 12 THE GID PARASITE AND ALLIED SPECIES. very completely and figured one accurately. Bloch very signifi- cantly adds that Leske numbered the parasites among the tape- worms, "Bandwiirmer," where, according to Bloch, they can not properly be reckoned, for reasons already given by him. It is evident from the last statement that Bloch had not overlooked Leske's Tsenia multiceps and that he believed lie was correcting an error by proposing the name Vermis vesicularis socialis. However, subsequent work on cestode life history has shown, the invalidity of all classifications which place vesicular worms in a group apart from the strobila forms and has justified Leske's judgment in uniting them. Unfortunately for Leske's name, Rudolphi (1810a) did not list it as a synonym of Ccenurus cerebralis, although he listed Leske's article in his bibliography. For this reason Leske's name has been very generally overlooked, as research in nomenclature has com- monly gone back through Rudolphi to the names quoted by him. Stiles and Stevenson (1905a) do not give Leske's name, Tsenia mul- ticeps, in their table of synonymy, and in selecting the oldest name available to them have overlooked the rather obscure references to Leske's unavailable article. On calling Doctor Stiles's attention to the omission he pointed out to me that Sherborn (1902a) refers to Leske (1780a) with the comment "No n. spp." I wrote Mr. Sherborn, asking him to verify this reference, which he very kindly did. In a personal communication he quotes substantially the part quoted by Kiichenmeister (1880a), and states that he over- looked the name in his former reading. Mr. Sherborn was also good enough to supply copies of Leske's illustrations. These show very close observation. Following the independent discoveries by Goeze and Leske of the cestode nature of the water bladder from the brain of giddy sheep, there arose some controversy as to which of them was entitled to priority. According to Braun (1894a), Boerner (1780a) published an article discussing this point and holding Goeze as the discoverer. Subsequently, Goeze (1782a) repudiated Boerner's article, deploring the misunderstanding between himself and Leske. He states that he has explained the situation in a previous publication, the date of which is not given and which is unavailable to me. Leske's priority is conceded by Rudolphi (1808a) and by Davaine (1860a). The matter of priority here is apparently not concerned in the nomen- clature, and what honor lies in priority of discovery belongs to Leske, so far as the available evidence shows. Goeze (1782a) divides his genus " Tsenia, Bandwurm," into two main classes as he calls them Tsenia visceralis, the visceral tape- worms, and Tsenia intestinalis, the intestinal tapeworms. Under the former he lists, among other species, ' ' Tsenia vesicularis cerebrina " HISTORICAL SKETCH OF MULTICEPS MULTICEPS. 13 from the brain of giddy sheep, Multiceps, the many-headed, with many heads and bodies in a common bladder. And later on he states that from the numerous heads one may call the parasite "Vielkopf (Multiceps)." From the above, Stiles and Stevenson (1905a) have taken the generic name Multiceps. The generic name used by Bloch (1780a) is evidently unavailable, being composed of two words and there- fore contrary to Article 8 of the International Code of Zoological Nomenclature, as given by Stiles (1905y): "A generic name must consist of a single word, simple or compound." Rudolphi (1809a) rejected Bloch's Vermis vesicularis as incon- gruous and unsystematic. Sherborn (1902a) is in error in listing Vermis Bloch 1782 as a generic name. The combination Vermis vesicularis is always used, whether with or without various specific names attached. As heretofore shown (p. 11), the earliest specific name of the parasite is that of Leske (1780a) as given in the name Tsenia multi- ceps. If the parasite in question is to be removed from the genus Tsenia, then the new combination must use the earliest available generic or subgeneric name, and since Goeze's (1782a) use of the scientific name Multiceps is evidently generic or subgeneric in intent, being clearly used to distinguish the many-headed gid parasite from the single-headed cysticercus forms, it is necessary to use it in the new name. The tendency for some time, and certainly a desirable tendency, has been to break up the large and heterogeneous group of animals formerly listed in the genus Tsenia, and to restrict the use of this name. The present situation has already been stated by Stiles (1905y) as follows: Most authors recognize that Tsenia is to be divided into the subgenera Tsenia, Multi- ceps (i. e. Cccnurus), and Echinococcus . Some authors, however, incline to recognize these subgenera as of full generic rank. It seems advisable to restrict the generic name Tsenia to those forms which have a cysticercus stage in the life history. These alone make up a large group with a fairly close similarity in the adult and larval stages. To retain in this already large genus forms having a ccenurus or echinococcus larva seems unnecessary and undesirable. Long ago Leuckart (1886d) wrote: The Coenurus * * * is related to the Cysticercus as a compound to a simple animal a sufficient reason for systematic zoologists to separate them. Generic rank is accorded to particular groups of species which in the course of evolution have attained distinctive characteristics, and I see no reason for withholding such rank from forms in which these distinctive characteristics occur in the larva instead of the adult. This point is of especial importance in a case of this sort 14 THE GID PARASITE AND ALLIED SPECIES. where the animal is found in the larval stage in the great majority of cases, the adult being seldom seen or recognized. This view is in accord with that of Stiles and Stevenson (1905a), from whom the following is quoted : Opinions may differ as to whether this group [Multiceps] should be given generic or subgeneric rank. Personally we see no serious argument against recognizing a distinct genus on basis of the "larval" stage. Adopting, then, the genus Multiceps Goeze, 1782a, and the species multiceps Leske, 1780a, as the oldest available names, the correct technical name of the gid parasite is Multiceps multiceps (Leske, 1780a), Hall, 1910/?. From 1782 to 1800, the latter date marking the beginning of Braun's (1894a) third period in helminthology, numerous observa- tions were made on gid, most of them merely confirming the previous work of Leske, Goeze, and Bloch, or adding minor points of more or less importance and interest. By 1800 the gid disease had been observed certainly for over a centu^ and a half and very likely for twenty-two centuries, its parasite had been named, described, and figured, and had a fairly large number of synonyms in addition to its correct name, the symptoms and pathology of the disease had been given, together with the symptoms of diseases simulating gid, and methods of operation had been used which only lacked aseptic pre- cautions to make them equivalent to good modern methods, and which were as good, perhaps, as most methods now in actual use. There remained, then, the work of finding out the life history and basing on that a rational prophylaxis. As a matter of fact the dis- covery of this life history by Kuchemneister and Von Siebold marks the beginning of the fourth and last period in helminthology. The contributions of the third period to the subject of gid are largely wrong and unnecessary theories of causation as well as unsatisfac- tory methods of treatment. In addition, the large amount of litera- ture in this period lists the parasite from several new hosts, often erroneously, and adds considerably to the synonyms by which it is known. During this period new records of the disease show a widening geographical distribution, and unsatisfactory and unsub- stantiated statements of its presence in the United States begin to appear as early as 1809. The essential contributions in the literature of this period have been covered hi tables and discussions to be given later, and the important events marking the modern period of helmin- thology may next be considered. Von Siebold (1844a) proposed as an explanation of "the true nature of bladderworms that they were cestode embryos which in attaining a new host had gone astray, ending as encysted, incompletely devel- oped forms. Thus Cysticercus fasciolaris of the mouse was held to be such an incomplete sexless modification of Tsenia crassicollis of the HISTORICAL SKETCH OF MULTlCEPS MULTICEPS. 15 cat. He ventured to predict that in time the various tapeworms would be identified in their relation to certain cysticercus, ccenurus, and echinococcus forms. Dujardin (1845a) advanced a similar theory, and this view or modifications of it became popular in scientific circles during the five or six years following Von Siebold's publication. It required the experimental work of Von Siebold and Kiichenmeister hi 1851 and 1852 to complete this half truth. In the meantime the advo- cates of spontaneous generation lost ground to those who urged that the bladderworms were altered, degenerate cestodes or were incom- pletely developed embryonal forms. A prominent champion of the last theory, Kiichenmeister (1851e), finally published a note stating that he had produced Tsenia cras- sitipes [= T. crassiceps] of the fox by feeding Cysticercus pisiformis. A little later (Kuchenmeister, 185 Id) he corrected this statement, changing his identification of the adult worm to T. serrata. This marks the beginning of the modern use of the now general experi- mental feeding methods of determining life histories. It remained for Von Siebold (1852a), the supporter of the theory of hydropic degeneration of bladderworms, to furnish additional proof that his theory was wrong, for this same year he produced the adult cestode from the gid bladderworm. The following year Kiichenmeister (1853e) succeeded in experi- mentally demonstrating, for the first time, the entire life history of a cestode. He fed Ccenurus cerebralis to a dog and produced a tape- worm which he called Tsenia coenurus. He then fed the gravid pro- glottids of this tapeworm to a sheep, and produced in it the early stages of the coenurus in the brain. From this experiment Kuchenmeister concludes that sheep are infected in pasture by dogs dropping proglottids. Other animals, he thinks, may also harbor the tapeworm, and he claims this would certainly be true of wolves in Hungary and Poland. This statement is evidently mere assertion, as it is not verified by the record of such a finding either at the time or subsequently. At this date no de- scription of T. cwnurus had been published and its anatomy had not been studied. Indeed, the following year Von Siebold (1854b) states that he finds the adult of C&nurus cerebralis to be Txnia serrata. While the occurrence of T. ccenurus in the wolf is a proba- bility, it is nothing more, so far as all available records show. On the evidence at hand Kiichenmeister formulated a set of rules for the prophylaxis of gid which is practically complete. It is as follows : 1. Feed dry food the year round and do not pasture. 2. Once or twice a year, purge the sheep and dogs in some inclosed place to get rid of tapeworms, and burn the feces. 16 THE GID PARASITE AND ALLIED SPECIES. 3. Do not, as is usually done, throw the heads of giddy sheep to the dogs> or, as Kiichenmeister after investigation finds to be done, throw the brain to the dogs before cooking the heads. Where there are wolves one must also bury or burn the intestines of those that are killed, and not throw them away to infect the fields. Such a program is not altogether practicable or necessary, but it only needs trifling amendment to bring it down to date. Had it been adhered to only as regards keeping dogs free from tapeworms and heads of giddy sheep away from carnivora for the last half cen- tury, gid would probably have been a rare disease by this, for it is really one of the most readily preventable of diseases. The next year Kuchenmeister's work was confirmed by Von Beneden (185458a Cattle ... .do do Sheep Germany Scultetus lf)72a.. Date of first certain case 1634. Cattle Germany (?) .... B runner 1 (>'.)!. According to Ktichenmeister Do " Berovla" Wepfer 1724a.. . . (1880a). "<;urtwillae" Sheep Germany Leske 1780a First recognized as a cestode para- Do do Goeze 1780a site. Independently recognized as a par- Do... Italy Fontana 1784a . . , asite. Cattle ..do do . Sheep Germany Sohrank 17SSa Cattle do . do Chamois. Alps Ketzius 179fla . A t least one case. Sheep England . Moorcroft 1792a Claimed to occur in France and Cattle do . .do... Italy also. African antelope.. . . Not given Rudolph! 1808a . Accepted here on basis of subse- quent findings. HOSTS AND OCCURRENCES OF MULTICEPS MULTICEPS. 31 List of certain or probable occurrences of the larval Multiceps multiceps Continued. Host. Locality. Authority. Notes and comments. Sheep France Bosc 181Ca Horse Bousset 1822a According to Gurlt (1831a) Do (') Hofacker 1823a Chamois. Paris, France De Blainville 1824a Sheep France Yvart 1827a Do do Dupuy 1831a.. cording to Braun (1894a). From spinal canal Horse Frenzel(Date?) According to Gurlt (1831a) and Do England Youatt 1834b Numan (1850b). St. Domingo goat. . . do Youatt 1836c symptoms given. Sheep. . . . Germany Pluskal 1844a. nurus on basis of subsequent findings. Do Austria do Do... Ireland Bellingham 1844a Goat Germany Klencke 1844a. Mouflon. . Montpelier, France Gervais 1847b One case Horse Ammon(?) According to Numan (1850b) Sheep (?) Storig(Date?). Found it twice in the medulla ob- Do Holland Numan 1850b longata, according to Numan (1850b). Angora goat do ....do in cerebrum, medulla oblongata, and in spinal cord. Coenurus is figured. Cattle Kempton, [Bavaria 1 *] Hering 1852a Occurred in 1850-51 Sheep Germany Hagmaier 1853a In spinal canal. Antelope Jacques and Lafosse Goat (?) 1854b. do Sheep Scotland . . McCall 1857a... Do France... Reynal 1857a. . Cattle .do.. do Sheep A If or t, France Valenciennes 1857a In spinal cord and brain' sent by Horse Vienna, Austria. Spinola 1858b Delafond. In spinal cord; specimen in veteri- Sheep Germany do.. nary school. In spinal cord. Goat Toulouse, France Baillet 1859b.. One certain and 1 possible infection Gazelle do . do of 4 experiment animals. Sheep Warschau . . . Leisering 1859a Eichler's subcutaneous specimen* Do Germany Leisering 18C2a found to be coenurus by Eichlerj Leisering, and Zenker. Gazelle (Antilope do do specimen; Eichler's specimen noted again. One case in a zoological park dorcap}. Horse Prussia Esse et al. 18G3a: Kei- One case; accepted on svmptoms Sheep Iceland per et al. 1804.a Krabbe 1864h and in view of other cases. Disease often seen here' accepted Cattle ..do do... on Krabbe's finding of adult worm in dogs. Rare; accepted as above Sheep Denmark do Claimed to occur; accepted as Cattle... . do ..do above. Do Do England Cooper 1805a.. Three cases. Chamois Germany Frauenfeld IStiSa Do. Sheep Vienna, Austria... . Bunion 1874a. . Several spinal cases seen by Roll. Sheep Germany. . . Miiller 1877a One case with coenurus in spinal Antelope (Bubalis sp.). Sheep Lyon, France South Australia Bertolus and Chau- veau 1879a. Dixon 1883a. cord; 1 in medulla oblongata. Host from Africa. Do Sardinia Parona 1884a Horse Culm, Germany Schwanefeld 1885a Contained one-sixteenth of a liter Sheep Buenos Ayres, Argen- Wemicke 1886a of fluid. nipnotragus egui- tine Republic. Germany L . Rabe 1889a. . Host from South Africa; in brain, nus (?). Sheep Montana, U. S Vet. Ed. Amer Sheep thyroid, lymph glands, and mus- culature. Do... New Zealand Breeder 1901 rand 3. Gilruth 1902a mortem findings. Goat Cape Colony Buckley 1904a Cow... ...do... Robinson 1905a... toms and post-mortem findings. One case. 32 THE GID PARASITE AND ALLIED SPECIES. List of certain or probable occurrences of the larval Multiceps multiceps Continued. Host. Locality. Authority. Notes and comments. Cattle . . France . Leblanc and Freger One case. Chamois Germany 1907a. Rothl907c Do. Cattle England Pollock 1908a Horse Shetland White 1909a. . One case; symptoms of gid and Cow Germany Pfab 1909a . . . recovery of parasite by operation. The only record found of the para- Sheep England Lloyd 1909a site from the vertebral canal in this host. Cattle Italy Vicariotto 1909a. Sheep . . . Montana and Wash- Hall 1909a . . Goat ington, D. C. Missouri, U. S Doctor Luckey in let- From personal correspondence with Cow - Germany ter of July 21, 1909. Borstelmann 1910a Doctor Luckey already noted. Bladderworm the size of pigeon egg Cattle do Pfab 19iav Bailly in 1861; claims that Prince has found this form in France. From Berlin Zoological Gardens. mus coy/rtw). Rabbit Italy Perroncito 1878h..., Listed from title; article not available. Squirrel (Sciurus England. . .. Cobbold 1879b Same case as Cobbold (18<>41>). vulpinust). Rabbit England (?) do Note that Alston has found cccnurus In Klippdas ( Ilyrax ca- Not given ....do rabbit. Error, due to confusing records of Ger- imi.ii.it. Gray squirrel United States Stewart 1880a. vais (1847a) and Pagenstecher (1877a). Probably a reference to Cobbold "s Rabbit do do (IHMb)case. Claimed to occur; no cases or authorities cited. OCCURRENCES OF MULTICEPS SERIALIS. 59 List of occurrences claimed for the larval Multiceps serial-is Continued. Host. Locality. Authority. Notes and comments. Squirrel (Sciurus France Cagny 1882a. One case. vulyaris). Rabbit do . do One specimen exhibited by Railliet in Do Italy Perroncito 1882a. . . discussion. Number of cases not given. Do Germany (?).. . . Braun 1885c One specimen. Do Russia (?) Reinitz 1885a Three specimens studied. Do France Railliet 1889n One case. Do ... .do Railliet 1889o Second spinal case; simultaneous con- Rabbit New Zealand Thomas 1889a nective tissue infection with 9 other coenuri. Not available; cited from Braun (1894a). Rabbit (Lepus cali- United States Brandegee 1890a . Manv cases in California; paper read in farnicus). Rabbit France Villain and Bascou 1882. Not available; cited from Morot (1900c); Do do 1890a. Leclerc 1890a one case. Not available; cited from Morot (1900c); Do .do Railliet 18911 several cases. One case; parasite lived over 2 years. Rabbit (L. varia- Russia Voigt 1891a . . One case. bilis). Rabbit (L. califor- United States. Curtice 1892g Number of cases not given; in Texas nicns and L. tcii- anus). Rabbit England Robinson 1892a and California. One case' scolices with 6 suckers. Do Do Italy Japan Condorelli-M a u g e r i 1893a. Janson 1893c One case; under pericardium. One case; listed as Ccenurus ccrebralis. Do France Megnin 1896. Not available; cited from Morot (1900c); Do do Lucet 1897b several cases. One case; 28 coBnuri. Do .do. Vignon 1897a . . . Not available; cited from Morot (1900c), Do United States. Ward 1897 b... who considers Vignon's Echinococciis a coenurus. Common in Nebraska. Horse .. .do Stiles 1898a Doubtful case, already noted under M. Rabbit... Italy Bosso 1898a . . . multiceps. One case. Rabbit (L. callotis). United States. Hassall 1898a. . . Specimens seen by Stiles or Hassall. Rabbit (L. cuni- Not North Amer- .do Do. culus). Rabbit . lea. France Railliet 1899b Specimen with many abnormal sco- Do .. .do Morot 1900c... lices. Four cases with 4, 11, 20, and 70 coenuri, Do.. .do.... Gallier 1900a. . in each host; 1 in eye orbit. One case. Do Siberia . Von Linstow 1901e.. Four specimens in St. Petersburg Do... Italy . Parona 1902f . . museum. Two cases. Do . France Buysson 1903a Rabbit (L. cuni- Germany (?) Kunsemiiller 1903a One case* specimen collected in 1874. c a lux domcsticus). Rabbit... England. ... Byerly 1905a . . One case. Do .. .do Jowett 19050. . . Has found it. Do. .. United States Ransom 1905d Specimen No. 1823 figured. Do. . . Scotland Taylor 1905a. Goat... India ... Gaiger 1907a. Do. Do... do Holterbach 1907a. Note of Gaiger's (1907a) case. Rabbit... Not given .do Sic. Cat .do .do . . Sic; error. Squirrel .do.... .do.... Sic. Sheep ...do.... ...do.... Sic; error. Horse .do .do Sic. Rabbit (L. califor- nicus). Rabbit United States ... .do S. E. Piper, in litt. Apr. 14. 1908. Curtice, in litt. July In Nevada; several coenuri fed to dog. In Colorado and California in 1887 and Goat India. .. 26, 1909. Dey 1909a 1888. One case: cysts In brain and connective Rabbit Switzerland Galli-Valerio 1909a tissue. Do England Gray 1909a Has seen it in eye orbit. Do. .. France Henry 1909a Cosnurus attained volume of 800 c. c. Do. New South Wales Johnston 1909a Listed Rabbit (Oryctolagus Victoria(?) Sweet 1909a. .. One case. cuniculus). Rabbit United States Dr. Young in litt. In North Dakota. "Sage rabbit".. . .do. Oct. 9, 1909. Thos. Large in litt In Idaho. Rabbit United States Jan. 6, 1910. Hall 1910/7 This article. 60 THE GID PARASITE AND, ALLIED SPECIES. The following specimens of M. serialis from the United States are available to the writer. Host. Locality. Collector and date. Collection. I.fpux californicus California Curtice 1890. B. A. I. coll. No. 1823. l.i pl ix sp. (?) (?) 1894 B. A. I. coll. No. 1826 Lepus callotis New Mexico Townsend 1896 B. A. I. coll. No. 2798. Lepus sp . . (?) (?) H A. I coll. No. 2G08 Do. . Michigan . Hayward 1904 B. A. I. coll. No. 3948 Lepus cali/ornicus California Adams 1905... B. A. I. coll. No. 3889. Lepus c. walla-walla Oregon Piper 1907 B. A. I. coll. No. 14728. Do. . do do B. A. I. coll. No. 14729. Do .do do B. A. I. coll. No. 14730. Lepus sp Nevada Hall 1910 B. A. I. coll. No. 15599. Do Nebraska Young 1905 Coll. Hall. The first of the above lists shows that Multiceps serialis has been claimed to occur in the hare, rabbit, squirrel, coypu, goat, horse, klippdachs, sheep, and cat. Records of its occurrence in the hare and rabbit are undoubtedly correct, the records from the squirrel are probably correct, those from the coypu and goat may be correct, the record from the horse is doubtful, as heretofore indicated, and those from the hyrax, sheep, and cat are errors. Cobbold (1864b) found a ccenurus in an American squirrel, Stiurus vulpinusl, which he thought might be the same species that Rose (1833a) found in "bladdery rabbits." This conclusion appears to be substantiated by the subsequent finding by Cagny (1882a) of a ccenurus in a squirrel, Sciurus vulgaris, which had been caught young and kept three years. Cagny's specimen was examined by Me'gnm ancl Railliet who pronounced it Coenurus serialis. Kunsemuller (1903a) thinks Cobbold's ccenurus may be C. serialis. If these authorities are right in their identification of this parasite, its rarity in this host is to be expected, as the squirrel's food is of such a nature, consisting as it does largely of nuts, that fecal contamination by carnivorous hosts of the adult worm would only occur very rarely. Stewart (1880a) writing from the United States, says: "The presence of this parasite [Ccenurus cerebralis} has been discovered in the liver of our gray squirrel and in rabbits, as well as in numerous sheep in this country." It is probable that the allusion to the para- site from the squirrel is a reference to Cobbold's (1864b) case of a ccenurus in an American squirrel. The reference to ccenurus forms having been found in American rabbits seems likely enough from our knowledge of the common occurrence of M. serialis in this country, but Stewart's record is uncertain, as he does not claim to have seen such a parasite, nor does he cite anyone who has. Lindemann (1867 a), according to a review by Rudnew (Linde- mann 1868b), described a Ccenurus lowzowi from the rabbit in Russia, in an article not available to the writer. This has since been very generally regarded as C. serialis. by helminthological writers, among DISCUSSION OF OCCURRENCES OF MULTICEPS SERIALIS. 61 whom are Pagenstecher (1877a), Moniez (1880a), Braun (1897a), and Kunsemiiller (1903a). The review of 1868 says there were no hooks in this form but other writers say the hooks were all the same size. Pagenstecher (187 7 a) says they were all the same size and finds the same thing in one scolex of his coenurus from Myopotamus coypus. Moniez (1880a) says the same and considers it either an error in observation or a teratological fact. Railliet (1899b) has found a great number of abnormalities in Multiceps serialis. In view of this fact and the unanimity of opinion concerning this form it has been accepted here as M. serialis. Pagenstecher (1877 a) describes a cosnurus which he identifies as Coznurus serialis from the neck of Myopotamus coypus. Reinitz (1885a) and Braun (1897a) think this form from the coypu is not M. serialis. Moniez (1880a) and Railliet (1882a) accept it as M. seri- alis, and Kunsemtiller (1903a) states that he agrees with Moniez and Railliet and disagrees with Reinitz and Braun. In view of this disagreement, the form is provisionally accepted as M. serialis, as originally described. Cobbold (1879b) has the following: The klipdas or dasse (Hyrax capensis) is infested by a tapeworm. * * * Under the name of Coenurus serialis a larval cestode has been described by Gervais, the same parasite being called Arhynchotxnia critica by Pagenstecher ("Zur Natur- geschichte der Cestoden." * * *). In the index this appears as " Coenurus serialis of the hyrax." Cobbold is in error in stating that Gervais described Coenurus serialis from the hyrax. As has been pointed out, his specimen was from the rabbit. Moniez (1880a) notes that Cobbold has confused Pagenstecher' s (1877a) statements, and Railliet (1882a) has stated that Cobbold has listed C. serialis from Ilyrax capensis as a result of some confusion. Gaiger's (1907^) and Dey's (1909oO records of M. serialis from the goat in India are provisionally accepted; a more extended discus- sion of these and other forms will be given in a subsequent paper dealing in part with the morphology of Multiceps spp. Holterbach's review of Gaiger's (1907/?) paper contains a number of errors in the list of hosts of M. serialis. The list of occurrences shows that the parasite has been reported from France, England, Scotland, Italy, Russia, Siberia, Switzerland, Australia, New Zealand, Japan, India, and the United States. Whether the parasite occurs in Germany is doubtful. Pagenstecher's (1877a) co3nurus was collected from a coypu in the Berlin Zoological Garden, and hence the origin of the parasite is in doubt. Reinitz (1885a) does not state where his three specimens were collected, but says that one was the specimen discussed by Braun (1883c) before the Dorpat Naturforscher Gesellschaft and the other two 62 THE GID PARASITE AND ALLIED SPECIES. were from Prof. Semmer. Braun (1883c) says of the specimen men- tioned that he owes it to "dem Herrn. stud. med. Hasenjager," from which it would appear that it was collected in Germany. Later, however, Braun (1897 a) lists the parasite from Russia on the authority of Reinitz (1885a) and Voigt (189 la), but in giving the distribution of this form he does not mention Germany. Still later, Braun (Braun u. Liihe, 1909a-), writing of the tapeworms of the domestic animals, refers to "Die in Deutschland noch nicht wohl aber in Frankreich beobachtete und sicher auch in Russland bei Hunden workom- menden T. serialis Baill." On the face of it, this statement can hardly be taken to mean more than that the adult T. serialis has not yet been observed in dogs in Germany, and Braun's English translator (Braun u. Liihe, 1910oO does not seem to have sufficient reason, especially as regards Germany for the statement that " T. serialis Baill. * * * occurs in dogs in France, and probably also in Russia, though not in Germany." Kunsemuller (1903a) does not give any locality for his specimens. The common occurrence of M. serialis in rabbits in the western part of the United States makes it unlikely that this parasite was imported into this country from the Old World, while its wide dis- tribution abroad and its apparent absence from the eastern part of this country makes it equally unlikely that it was carried abroad from this country. Its presence in Oregon and in Siberia points to the strong possibility of its having spread by way of far northern routes over its present wide range of distribution. M. serialis has been recorded from the vertebral canal by Leblond (1837a) and Railliet (1889o), in the latter case with an accompanying infection of the more usual connective-tissue locations. It has been recorded from the pericardium once by Condorelli-Maugeri (1893a), from the eyelid by Byerly (1905^), and from the orbit of the eye by Gray (1909^), and by Mr. S. E. Piper of the Bureau of Biological Survey of the Department of Agriculture in data furnished the writer. The number of parasites varies from one, a very common record, to 70 in one case of Morot (1900c), and in size the cyst may attain a volume of 800 c. c., as in the case of Henry (1909 A-). The parasite may live over two years according to Railliet (1891i). Abnormal specimens have been noted by Pagenstecher (1877a) from the coypu, by Robinson (1892a), Railliet (1899b), and Galli-Valerio (1909ar), from the rabbit, and Lindemann's (1867a) specimen was probably such. Successful operations for the parasite have been noted by Railliet (1889n) and Byerly (1905^). Mr. Piper, who has furnished the Bureau collection with speci- mens as noted above, has also furnished us data stating that the OCCURRENCES OF ADULT MULTICEPS SERIALIS. 63 parasite was found in 7 out of 12 rabbits examined in Oregon, a pint of cysts being taken from the peritoneal cavity of one. Mr. Piper also collected M. serialis in Nevada in 1908, as noted in the table, and fed a number to a dog. The dog was shipped to this laboratory, but did not develop the adult parasite, probably owing to diarrhea resulting from intestinal irritation by too many scolices. The writer has since collected M. serialis in Nevada, and developed the adult worm by feeding scolices to a dog. Mr. Graybill, of this laboratory, has also collected M. serialis in Texas and fed it to a dog. Doctor Young, of the University of North Dakota, writes under date of Octo- ber 9, 1909, that there is a specimen in the university collection, unlabeled, and that rabbits which appear to be infected are seen in North Dakota; he himself has seen such a rabbit. Doctor Shantz, of the Bureau of Plant Industry, has seen such rabbits in Kansas and Colorado, and Mr. E. F. Chilcott of the same Bureau says they are common in South Dakota. Kaupp's (19Wa) statement that M. serialis is not common in the United States is hardly accurate. In certain Western States it is very common. The occurrence of the larval parasite in the muscles of its host, especially in the leg muscles, a common site, and its occurrence in such relatively enormous sizes, numbers, or quantities as are given hi the more extreme cases of Henry (1909^), Morot (1900c), and Mr. Piper, may be looked upon as an adaptation favorable to the parasite, serving to impede the locomotion of the secondary host and so increase the likelihood of its being captured by some carnivore which may serve as the primary host of the parasite. Brandegee (1900a) has also pointed out the presence of an adaptation here. THE OCCURRENCES OF THE ADULT MULTICEPS SERIALIS. The dog is the only host in which the adult Multiceps serialis has been found or produced. Thomas's (1889a) attempts to infect cats and ferrets by feeding them the larval cestodes failed, according to Braun (1894a), and a surmise such as that of Brandegee (1890a) that the wolf, coyote, lynx, and fox may act as hosts, has, of course, only the value of a surmise. At the same time, Baillet (1866b) early called attention to the fact that the larval parasite was found in the wild rabbit more commonly than in the domestic rabbit, and sur- mised that the usual host was some wild carnivore. Galli-Valerio (1909aO failed to develop the adult worm on ingesting two living heads from the larval parasite. The writer also has similarly failed to develop the adult worm on ingesting three living heads from the larval parasite. 64 THE GID PARASITE AND ALLIED SPECIES. List of occurrences of the adult Multiceps serialis in the dog. Locality. Authority. Notes and comments. France Baillet 1858b... By experiment. Do... Baillet 1863a Do. Do Baillet 1866b.. Found several times. Do Bertolus and Chauveau One case in a cosmopolitan "dog of the regiment." Italy 1879a. Perroncito 1878a Not available; based on Railliet 's (1882a) state- Do Perroncito 1882a ment that Perroncito failed to infect sheep from Canwus serialis. By experiment. New Zealand Thomas 1889a. Not available* cited from Braun (1894a) - by ex- France Neumann 1892a .. periment. Several times. Japan . Janson 1893c By experiment. France Railliet 1893a By experiment; claimed that Neumann has also North America Ward 1895b produced it; I can not verify claim. Listed. Do Sommer 1896c Stated on the authority of Stiles. United States. Ward 1897b One case out of 20 dogs in Nebraska* others im- Do Stiles 1898a plied. Parasite seen by Stiles. Do Stevenson 1904b Two cases out of 35 dogs in Nebraska; 20 speci- Australia Cobb 1905a mens. One specimen; identification not positive. United States. Ransom 1905d Specimen No. 2839 figured. India Gaiger 1907a By experiment. New South Wales. Johnston 1909a . . Rare. United States Hall 1910/J This article. ECONOMIC IMPORTANCE. As has been stated, Multiceps serialis is of comparatively little economic importance. It deserves attention from an economic stand- point largely because some scientists, especially the Italian, insist on identifying it with the highly important M. multiceps. Rose (1833a) states, as before mentioned, that when warreners meet with infested rabbits they puncture the bladder, squeeze out the fluid and send the animal to market. According to Martel (1909or), this custom of puncturing through the skin of infected rab- bits is still in vogue in France. While the idea of eating the parasite is not a pleasing one, the danger from doing so is negligible as the parasite is apparently not transmissible to man,, as Galli-Valerio's (1909or) and the writer's experiments along this line indicate. Moreau (1909r) in a note on abattoir inspection in France, lists muscular coe- nurosis of hares and rabbits as sufficient cause for total condemnation of the carcass, but probably this practice would only be followed in such cases as those listed by Morot (1900c), where rabbits were condemned owing to infestation with 11, 20, and 70 cosnuri each. In Morot's cases, a rabbit infested with only 4 ccenuri was returned for food after the removal of the diseased parts. The writer finds that in the western United States the carcasses of rabbits infected with M. serialis are thrown away as unfit for food. SYNONYMY OF MULTICEPS SEEIALIS. 65 SYNONYMY. The generic synonymy has already been given under Multiceps multiceps. Species MULTICEPS SERIALIS (Gervais 18473) Stiles and Stevenson 19053. 1828. E [chinococcus] veterinorum(t) of de Blainville 1828a; misdetermination . 1833. Ccenurus cerebralis Lamarck and Rudolph! of Rose 1833a; this combination should be attributed to Rudolphi 1808a; error; misdetermination. 1837. Csenurus cerebralis of Leblond 1837a; error; misdetermination. 1844. Ccenurus cerebralis of Rose 1844a; misdetermination. 1847. Ccenurus serialis Gervais 1847a. 1855. Ccenurus serdalis Gervais of Goldberg 1855a; in synonomy of Tsenia ccenurus; this combination should be attributed to Goldberg 1855a; misprint. 1863. Tsenia serialis (Gervais 1847a) Baillet 1863a; first naming of strobila form. 1863. Ccenurus cerebralis? leporis cuniculi Baillet of Diesing 1863b; in synonomy of Tsenia ccenurus; not at present available, cited from Diesing 1864a, identical; this combination should be attributed to Diesing 1863b. 1863. Tsenia ccenuri cuniculi Baillet of Diesing 1863b; in synonomy of Tsenia ccenurus; this combination should be attributed to Diesing 1863b. 1864. Ccenurus cuniculi (Diesing 1863b) Cobbold 1864b; name taken from MSS. of Rose. 1867. Ccenurus lowzowi Lindemann 1867a; not available, cited from Lindemann 1868b; same form used once by Braun 1894a. 1868. Tsenia ccenurus of Cobbold 1867a; error. 1877. Ccenurus loivtzowi Lindemann of Pagenstecher 1877a; this combination should be attributed to Pagenstecher 1877a; misspelling. 1877. Ccenurus nov. spec, of Pagenstecher 1877a; Pagenstecher refers thus to the form which he identifies as Ccenurus serialis. 1877. Ccenurus serialis Gervais of Davaine 1877a; this combination should be attrib- uted to Davaine 1877a; misspelling. 1877. Tseniaserialis Baillet of Davaine 1877a; space omitted. 1879. Arhynchotsenia critica Pagenstecher of Cobbold 1879b; error. 1882. Caenurus serialis (Gervais 1847a) Perroncito 1882a. 1882. Csenurus saerialis Gervais of Perroncito 1882a; this combination should be attrib- uted to Perroncito 1882a; misspelling. 1882. Ccenurus serialis Baillet of Ziirn 1882