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Oh) ‘hance avs Oi WA Nap nh Mgh oh Ally Simyry Ne) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. XXXV EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES, AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN—SMITHSON ————— ——,sonian Instiz, yx, XO (No. 1740) AUG 16 917 + Vo 0 ; yor hi ‘Onal M use / if CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION Ibe aes) - peo SE MEN. THis volume forms the thirty-fifth of a series, composed of original memoirs on different branches of knowledge, published at the expense and under the direction of the Smithsonian Institution. The publication of this series forms part of a general plan adopted for carrying into effect the benevo- lent intentions of James Smiruson, Hsq., of England. This gentleman left his property in trust to the United States of America to found at Washington an institution which should bear his own name and have for its objects the ‘increase and diffusion of knowledge among men.’’ This trust was accepted by the Government of the United States, and acts of Congress were passed August 10, 1846, and March 12, 1894, constituting the President, the Vice- President, the Chief Justice of the United States, and the heads of Executive Departments an establishment under the name of the ‘‘ Smirmsontan Insrtr- TUTION, FOR THE INCREASE AND DIFFUSION OF KNOWLEDGE AMONG MEN.’’ The members of this establishment may hold stated and special meetings for the supervision of the affairs of the Institution and for the advice and instruction of a Board of Regents to whom the financial and other affairs are intrusted. The Board of Regents consists of two members ex-officio of the establish- ment, namely, the Vice-President of the United States and the Chief Justice of the United States, together with twelve other members, three of whom are appointed from the Senate by its President, three from the House of Repre- sentatives by the Speaker, and six persons appointed by a joint resolution of both Houses. To this board is given the power of electing a Secretary and other officers for conducting the active operations of the Institution. To carry into effect the purposes of the testator, the olan of organization should evidently embrace two objects; one, the increase of knowledge by the addition of new truths to the existing stock; the other, the diffusion of knowl- edge, thus increased, among men. No restriction is made in favor of any kind of knowledge, and hence each branch is entitled to and should receive a share of attention. The act of Congress establishing the Institution directs, as a part of the plan of organization, the formation of a library, a museum, and a gallery of art, together with provisions for physical research and popular lectures, while it leaves to the Regents the power of adopting such other parts of an organiza- tion as they may deem best suited to promote the objects of the bequest. Ill IV ADVERTISEMENT After much deliberation, the Regents resolved to apportion the annual income specifically among the different objects and operations of the Institution in such manner as may, in the judgment of the Regents, be necessary and proper for each, according to its intrinsic importance, and a compliance in good faith with the law. The following are the details of the two parts of the general plan of organi- zation provisionally adopted at the meeting of the Regents December 8, 1847: DETAILS OF THE, FIRST PART (OF TES Rane I. To rycrease KNowrepcr.—It is proposed to stimulate research by offering rewards for original memoirs on all subjects of mvestigation. 1. The memoirs thus obtained to be published in a series of volumes, in a quarto form, and entitled ‘‘ Smithsonian Contributions to Knowledge.’’ 2. No memoir on subjects of physical science to be accepted for publication which does not furnish a positive addition to human knowledge, resting on original research; and all unverified speculations to be rejected. 3. Each memoir presented to the Institution to be submitted for examina- tion to a commission of persons of reputation for learning in the branch to which the memoir pertains, and to be accepted for publication only in case the report of this commission is favorable. 4, The commission to be chosen by the officers of the Institution, and the name of the author, as far as practicable, concealed, unless a favorable decision be made. 5. The volumes of the memoirs to be exchanged for tlie transactions of literary and scientific societies, and copies to be given to all the colleges and principal libraries in this country. One part of the remaining copies may be offered for sale, and the other carefully preserved to form complete sets of the work to supply the demand from new institutions. 6. An abstract, or popular account, of the contents of these memoirs to be given to the public through the annual report of the Regents to Congress. If. To increase Knowxiepcr.—lt is also proposed to appropriate a portion of the income annually to special objects of research, under the direction of suitable persons. 1. The objects and the amount appropriated to be recommended by coun- sellors of the Institution. 2. Appropriations in different years to different objects, so that in course of time each branch of knowledge may receive a share. ADVERTISEMENT Vv 3. The results obtained from these appropriations to be published, with the memoirs before mentioned, in the volumes of the Smithsonian Contributions to Knowledge. 4, Examples of objects for which appropriations may be made: (1) System of extended meteorological observations for solving the prob- lem of American storms. (2) Explorations in descriptive natural history, and geological, mathe- matical, and topographical surveys, to collect material for the formation of a physical atlas of the United States. (8) Solution of experimental problems, such as a new determination of the weight of the earth, of the velocity of electricity, and of light; chemical analyses of soils and plants; collection and publication of scientific facts, accu- mulated in the offices of Government. (4) Institution of statistical inquiries with reference to physical, moral, and political subjects. (5) Historical researches and accurate surveys of places celebrated in American history. (6) Ethnological researches, particularly with reference to the different races of men in North America; also explorations and accurate surveys of the mounds and other remains of the ancient people of our country. I. To pirruse Knowiepcs.—I/t is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge not strictly professional. 1. Some of these reports may be published annually, others at longer in- tervals, as the income of the Institution or the changes in the branches of knowledge may indicate. 2. The reports are to be prepared by collaborators eminent in the different branches of knowledge. 3. Each collaborator to be furnished with the journals and publications, domestic and foreign, necessary to the compilation of his report; to be paid a certain sum for his labors, and to be named on the title-page of the report. 4. The reports to be published in separate parts, so that persons interested im a particular branch can procure the parts relating to it without purchasing the whole. 5. These reports may be presented to Congress for partial distribution, the remaining copies to be given to literary and scientific institutions and sold to individuals for a moderate price. VI ADVERTISEMENT The following are some of the subjects which may be embraced in the reports: I. PHYSICAL CLASS. 1. Physics, including astronomy, natural philosophy, chemistry, and meteorology. 2. Natural history, including botany, zoology, geology, etc. 3. Agriculture. 4. Application of science to arts. II. MORAL AND POLITICAL CLASS. 5. Ethnology, including particular history, comparative philology, antiq- uities, ete. 6. Statistics and political economy. 7. Mental and moral philosophy. 8. A survey of the political events of the world; penal reform, ete. Ill. LITERATURE AND THE FINE ARTS. 9. Modern literature. 10. The fine arts, and their application to the useful arts. 11. Bibliography. 12. Obituary notices of distinguished individuals. II. To pirruse Knowrepce.—Ilt is proposed to publish occasionally separate treatises on subjects of general interest. 1. These treatises may occasionally consist of valuable memoirs translated from foreign languages, or of articles prepared under the direction of the Institution, or procured by offering premiums for the best exposition of a given subject. 2. The treatises to be submitted to a commission of competent judges previous to their publication. ADVERTISEMENT VII DETAILS OF THE SECOND PART OF THE PLAN OF ORGANIZATION. This part contemplates the formation of a library, a museum, and a gallery of art. 1. To carry out the plan before described a library will be required con- sisting, first, of a complete collection of the transactions and proceedings of all the learned societies of the world; second, of the more important current period- ical publications and other works necessary in preparing the periodical reports. 2. The Institution should make special collections particularly of objects to illustrate and verify its own publications; also a collection of instruments of research in all branches of experimental science. 3. With reference to the collection of books other than those mentioned above, catalogues of all the different libraries in the United States should be procured, in order that the valuable books first purchased may be such as are not to be found elsewhere in the United States. 4. Also catalogues of memoirs and of books in foreign libraries and other materials should be collected, for rendering the Institution a center of biblio- graphical knowledge, whence the student may be directed to any work which he may require. 5. It is believed that the collections in natural history will increase by donation as rapidly as the income of the Institution can make provision for their reception, and therefore it will seldom be necessary to purchase any article of this kind. 6. Attempts should be made to procure for the gallery of art casts of the most celebrated articles of ancient and modern sculpture. 7. The arts may be encouraged by providing a room, free of expense, for the exhibition of the objects of the Art Union and other similar societies. 8. A small appropriation should annually be made for models of antiqui- ties, such as those of the remains of ancient temples, etc. 9. The Secretary and his assistants, during the session of Congress, will be required to illustrate new discoveries in science and to exhibit new objects of art. Distinguished individuals should also be invited to give lectures on sub- jects of general interest. In accordance with the rules adopted in the programme of organization, each memoir in this volume has been favorably reported on by a commission appointed for its examination. It is, however, impossible, in most cases, to verify the statements of an author, and therefore neither the commission nor the Institution can be responsible for more than the general character of a memoir. OFFICERS OFf THE SMITHSONIAN INSTITUTION. WOODROW WILSON, PRESIDENT OF THE UNITED STATES, EX OFFICIO PRESIDING OFFICER OF THE INSTITUTION. EDWARD DOUGLASS WHITE, CHIBF JUSTICE OF THE UNITED STATES, CHANCELLOR OF THE INSTITUTION. CHARLES D. WALCOTT, SECRETARY OF THE INSTITUTION. RICHARD RATHBUN, ASSISTANT SECRETARY. VIII MEMBERS EX OFFICIO OF THE INSTITUTION. NVOODR Owen VAIS ONGamIsEte IG einriciele cele sicieiele seus President of the United States. HELONUAG UN e a MUARSIDATI ec. 2 costs 6 secre aoe o 3d Vice-President of the United States. Hpwarp WDOUGKASS§ WHITH. ..c... 6 e.s sess Chief Justice of the United States. ROBERT LANSING. ...--.-.2-00. +020 se seen Secretary of State. Wituram Gisss McApoo............ wee... Secretary of the Treasury. Nanarony ]Dinorene, JBVNae)5 6 gone aaue ooo aineOODe Secretary of War. TETONTAG HAN (ACIS CHRING ORY) .) 01a iiacic a6 2 sce 302 Attorney-General. ALBERT SIDNEY BURLESON...........:..-..- Postmaster-General. Josmmnmias IDA do's coos colon Oe Goi ane Secretary of the Navy. IDA END IRTSIEGE IAG, gene oe one eo ome Secretary of the Interior. DAWA HIRAWE KIN OUSTON se <2) 2 ete) ella Secretary of Agriculture. \WWwinnanut (Cione Jissinyanat), pg oaoao serosa ooT Secretary wy Commerce. Wiiiam BaucHop WILson................ Secretary of Labor. REGENTS. Epwarp Doucuass WHITE Tomas R. MarsHanu Henry Casor Lopen......:..... WAnTseTAUVire) ica TOMBE: se tee ee teks Henry Frenco Ho3uis......... Scour (WERRIS. g 2 hry btae le easels Afavintspl be IG wOhanowd acidinde 40050008 HENEST Whe ROBERTS ss os ene Anprew D. WHITE Citizen ALEXANDER GraHAM BELL....... Citizen GEORGE GRAY Citizen CHarnEs HY. CHOATE, JR... 42.00 Citizen Joun B. Henperson, JR Citizen Cuarutes W. F'AarrBANKS Citizen x Member Member Member Member Member Chief Justice of the United States, Chancellor. Vice-President of the United States. of the Senate. of the Senate. of the Senate. of the House of Representatives. of the House of Representatives. Member of the House of Representatives. of New York. of Washington, D. C. of Delaware. of Massachusetts. of Washington, D. C. of Indiana. CONTENTS. AUTSTER ERED: 2 boi Ho SSS Goad OID OO ania Ieee teicher choi eer cicicici WSistore OlicerswVMembers ANd UREPENtS a5. asics sale Sees Weegee does Hanes aee yee dee vill ArtTIcLE 1 (1718). The Young of the Crayfishes Astacus and Cambarus. By E. A. ANDREWs. Published 1907. 4to, 79 pp., 10 pls. ARTICLE 2 (1723). The Apodous Holothurians. A Monograph of the Synaptidae and Mol- padiidae, including a Report on the Representatives of these Families in the Collections of the United States National Museum. By Hupert Lyman Ciark. Published 1907. 4to, 251 pp., 13 pls. ARTICLE 3 (2382). A Contribution to the Comparative Histology of the Femur. By J. 8. Foorr, M. D., edited by ALES HrpoxiéKa. Published 1916. ix, 242 pp., 38 pls. XI Fs a 7 ee 4 a a - ¥ is ‘ Le “ sc Roh: - Zs it , Fapes' = \ “ Kn ty” 97 io x a 540) pat ae, -- ~ ia. - 5 We. ie oe 3 lg Bs hes tT: SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE PART OF VOLUME XXXV ‘| THE YOUNG OF THE CRAYFISHES = ASTACUS AND CAMBARUS BY E. A. ANDREWS ae RHE” Re SviFEVSo, Ea » oe Ane oe N (No. 1715) : ‘ A CITY OF WASHINGTON re: PUBLISHED BY THE SMITHSONIAN INSTITUTION \\ pil th 1907 . | ! | On a ee i TR, Pe ee yh eee nti om S - ? 7 - : y SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE PART OF VOLUME XXXV fas 1OUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS BY E. A. ANDREWS (No. 1718) CITY OF WASHINGTON - PUBLISHED BY THE SMITHSONIAN INSTITUTION 1907 Commission to whom this memoir has been referred : WALTER FAXON WILLIAM KEITH BROOKS WILLIAM PERRY HAY WASHINGTON, D. (3 PRESS OF JUDD & DETWEILER, INC, 1907 ADVERTISEMENT. The present memoir by Professor E. A. Andrews, of the Johns Hopkins University, on ‘‘The Young of the Crayfishes Astacus and Cambarus,’’ forms part of Volume XXXV of the Smithsonian Contributions to Knowledge. The memoir describes and illustrates the young of two kinds of crayfishes, one from Oregon and one from Maryland, which represent the two most diverse forms found in North America. Of these, one genus, found only in North America, is widely distributed all over the United States, except the Pa- cific Slope; while the other is restricted almost exclusively to the Pacifie Slope in North America, and at the same time it is almost the only genus in Europe and Asia. This memoir fills a gap in the knowledge of these common animals that still remained notwithstanding the extensive researches of Huxley and many others. It determines the form and habits of the first, second, and third larval stages; gives the first detailed description and illustrations of the appendages of the first and second stages; describes the hitherto unknown nature of sue- cessive mechanical attachments of the offspring to the parent; and opens up the problem of the nature and causes of the incipient family life in the eray- fish. The new facts and comparisons add to the data for solution of the impor- tant problems of the geographical distribution and the origin of the species of crayfish, and they furnish a basis for practical application to the problems of artificial culture and introduction of new kinds of crayfish. In accordance with the rule adopted by the Smithsonian Institution, the work has been submitted for examination to a commission consisting of Dr. Walter Faxon, of the Museum of Comparative Zodlogy, at Cambridge, Mas- sachusetts; Prof. W. K. Brooks, of the Johns Hopkins University, and Prof. W. P. Hay, of Howard University, who recommended its publication in the pres- ent series. Cuas. D. Watcort, Secretary. SMITHSONIAN INSTITUTION, Wasutneron, June, 1907. " CONTENTS. Ti, TimiiaglinciiiGny: oats eco o cee o oO OReIGRS OO OORt ne ee ene Inne ieee ree pe Seema On Til, TENSiORIGEM wc bk eB ocdis Oe Gee cee Seber RRR eRe ee CR Cee cee ETS MI EReAIST ACIS MICTIVUSCUIUS) vec che ace cies ee ae + Seine 2: Neiese er eterna Sys opel cee late eels FMM OCHS OL ree Reese creel tay= ya Sie eh ss 2slovar ice 76 one. Sis iloyau sote perch sal o abe yeeorass/aevore aver SMRS OLIRCOMOLMRITI ALDI All cycee init eye e irae GO Oe ae aaa EIR Ona e com concer res Da tco ol Diac Geihincdlarvallstace= Sizes habits); tropisms <2 .272---.--2 3-2. +---- = - 7. Fourth and later larval stages: Number of moults and rates of growth .-. &, IRA Or GGse kOaell iminiainys 6 oooeecoboeenoooeeDDoacooUesdG56 ra dete Wan Compamsouseanduconelusionseacs. co) 42- 2 ee: ts) see ee oe eee le Page ¥ 9 10 11 12 14 41 41 41 42 43 The Young of the Crayfishes Astacus and Cambarus. By E. A. ANDREWS. INTRODUCTION. The object of the present paper is to figure and describe the external forms and the appendages of the early larval stages of the crayfishes Astacus and Cambarus and to illustrate the details of the connections that exist be- tween these larve and the mother. It is well known that the genus Cambarus is found only in North America and here only to the east of the Rocky Mountains, while the genus Astacus is the only one found in Europe and Asia and in America is found almost ex- clusively west of the Rocky Mountains. It is thus natural that the history of scientific knowledge of crayfishes has been first the study of Astacus in Europe and later the study of Cambarus in the eastern United States. The Astacus of the Pacifie States remains less well known. Despite all the work that has been done upon these common animals, several parts of their life histories have received scant attention. The geo- graphical distribution and systematic description have been studied in detail by Hagen, Faxon, Huxley, Ortmann, and others; the embryology minutely ob- served by Rathke and by Reichenbach; and the general knowledge of crayfish natural history added to again and again since the days of Roesel von Rosen- hof. Yet little attention has been given to the study of the young crayfish after it leaves the egg; a comparative neglect that naturally arose from the centering of scientific interest upon the larval changes of marine crustacea in which re- markable metamorphoses oceur. When these metamorphoses were established by Vaughn Thompson and others it was already known from the work of Rathke that the crayfish hatches from the egg in essentially the adult shape and thus passes through no series of metamorphoses. Interest in the crayfish young was then restricted to the fact that it was exceptional in having no metamorphosis. In the preoccupation of students of crustacean life histories in study of metamorphoses the young of the crayfish were left without any illustrations excepting only those given by Rathke to show the condition of the embryo when nearly ready to hatch and the two wood euts given by Huxley. These latter illustrations were evidently 5 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS made from specimens preserved in alcohol, and, however excellent for their pur- pose, fail to give the just proportions of the larve, since all alcoholic specimens of young crayfish are much swollen and distorted. While two more figures of the young erayfish have recently been given, (Andrews, ’04) details of external structure are still lacking. Having obtained the young of both Astacus and Cambarus by hatching the eges in the laboratory, it seemed well to fill in some of the gap in our knowledge of the early larval life of crayfish, and especially so as the details of a curious mode of connection with the parent were here first made evident. HISTORICAL. A brief statement of the history of our knowledge of the young of erayfish will show that as yet the character of the appendages in the early stages, the exact number of stages present in the life of the young while associated with the mother, and the nature of the means by which the young are held attached to the mother have waited discovery and illustration. To Roesel von Rosenhof belongs the credit of an enthusiastic appreciation of the care of the mother crayfish for the young; the observation that the young are transparent and like the parent; the description of their crowding upon the abdomen of the parent and of their finally forsaking her after a few days, during which, however, they would return to her at times as if reealled by a signal. Rathke (’29) was chiefly concerned with the embryology of the crayfish, but also described something of the growth in proportions and in internal anatomy of the young after hatching and gave figures of the embryo and of some of its appendages shortly before hatching. Reichenbach (’86) also added to his classie study of the embryology only a figure of the abdomen of a recently hatched larva. Soubeiran (’65) measured young crayfish grown at the farm of Clairfon- taine and recorded their moultings and rates of growth: More facts of this same kind were gathered by Chantran (’70, ’71) from prolonged study of larve reared in the laboratory of M. Coste. Chantran also discovered a peculiar fila- ment that held the young to the egg-shell after hatching and he finally convinced himself that the young ate their egg shells and their cast-off skins. Some other observations upon the size and times of moulting of young erayfish in Sweden were also made by Steffenberg (’72). Huxley (779) illustrated the recently hatched larve by two wood euts, one showing the early larva fastened to the maternal pleopods by its pecuharly re- eurved elaws, which he first described and figured, and the other a dorsal view of such a larva. He rectified the previous statement that the young at hateh- ing are exactly like the adult and pointed out their differences in lack of sete, THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 9 in lack of abdominal appendages on the first and sixth somites, and in various proportions. _ In America Hagen (’70) in his fundamental study of our crayfish observed the young of Cambarus attached to the mother in aleoholic specimens and re- corded the dimensions of young in which the sexes were externally recogniz- able. From like specimens Faxon (’85) in his revision of American crayfish was able to make a number of comparisons with specimens of young Astacus pallipes from France, and also to demonstrate the important facts that the young Cambarus agreed, in general, with Astacus as deseribed by Huxley, but that in the early larva as in the adult, Cambarus showed no vestige of the gill found on the last thoracic somite in Astacus. These facts were verified by Steele (’02), who for the first time described living larve of Cambarus hatched from eggs in the laboratory and also recorded facts as to the habits and sizes of young larve. From like living material Andrews (’04) added details of hatching and behavior of these larve with figures of side views of living larve in the first and in the second stages and noted the occurrence of two successive attachments of the larve by means of special structures. To complete that last preliminary paper by adding details and many new facts observed in Cambarus and to deseribe similar stages in our American Astacus is the purpose of the present paper. II. ASTACUS LENIUSCULUS. As far as known, no observations have been made upon the life histories and habits of any American crayfish of the genus Astacus and as the following faets were gained from animals kept in Baltimore far from their native habitat they will need the corroboration of future studies made in the Pacific States, but for the present they supply all our knowledge of the young of American Astacus. The material for study of the young Astacus was obtained as follows: Sixty- four specimens were received February 23, 19094, by express from Portland, Oregon, and, though packed only in wet excelsior, ten females and eighteen males survived the journey and lived in running water in the laboratory for some time. Shipped without selection of sex, the sixty-four were found to be made up of thirty-one females and thirty-three males, which indicates an equal distribu- tion of the sexes. The males, however, seem to have endured the journey better than did the females. During Mareh, April, May, and June these crayfish died slowly one by one, leaving one survivor in October. As they were fed from time to time with small oligochxte, with fragments of crayfish and with pieces of frogs, the cause of the very slow and lingering deaths was not evident, but probably the food was not sufficient for such large and active crayfish. 2 10 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS It was decided that this Astacus was probably Astacus leniusculus Dana as distinguished from 4. Trowbridgii by Faxon (’85) and the following observa- tions upon their habits in captivity seem to be all that is known of this species, beyond its specifie characters. The largest specimen was a female measuring 155 mm. from tip of rostrum to end of telson, 205 mm. from tip of chela to tip of telson, 40 mm. wide across the thorax and 35 mm. deep; the abdomen was 45 mm. wide and its tail fan expanded to a width of 70 mm., exclusive of the sete. With only about one-half of its eggs left upon its pleopods this female weighed 106 grams. The smooth clean shells and large well colored claws gave these crayfish an attractive and decidedly lobster-like appearance, which was enhanced by their very active pugnacious nature. In shallow water they quickly responded to approaching objects and readily threw themselves into a defensive attitude, leaning back with the whole anterior region raised high from the bottom of the tank, thrusting their brilliantly colored claws above them high into the air and either holding them wide apart and open or clashing them together towards an approaching object, towards which they lunged, or even seemed to spring. Their quickness to react to distant objects, their quick reflexes and irrita- bility led me to suppose they might well be carnivorous and their clear colors suggested a life in clear water, but the collector reported that the Willamette River, whence they came, was a rather muddy stream, though formerly used as water supply for Portland. In captivity they were nocturnal, lying quiet and away from the lighter parts of the tank in the daytime and crawling about in the night. They sometimes injured one another and also ate parts of their dead fellow-crayfish and, as is so common with many kinds of crayfish, there were cases of regeneration, which may have followed from injuries caused by their -pugnacious and carnivorous habits. ‘One malé 95 mm. long, when received, had two regenerating limbs. The left chela was represented by its original basal two segments and by a delicate new limb only 9 mm. long protruding from the truncated end of the second large segment. ‘This little, bluish protuberance was made of six movable segments and it was movable upon the old limb whence it sprang, so that to make the seven normal movable segments of a complete limb a reduction in the number of movable joints must take place, probably by the base of the new growth ceasing to move upon the old second segment. The other regenerating limb was the third left walking leg and here again the old second segment bore a soft protuberance, which, however, was as yet not segmented. Thus in this Astaeus, as is common in other crayfish, re- generations had started from the preformed breaking plane, and in handling this same specimen, though dead, the right third walking leg fell off at the breaking plane. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 11 In captivity shedding, or eedysis, took place in early summer, in some; thus on June 11, when the water had risen from 9° C. in March to 22° C., one large male cast its shell as one perfect piece containing also the ‘‘teeth’’ and other part of the lining of the ‘‘stomach.’’ The creature was then quite soft and easily indented by touch upon its carapace; it lay inert but could be made to crawl in a somewhat palsied way and even to flap its abdomen. On each side of the heart region the carapace bore a white, indented, scar, as if made by the claws of some other crayfish, so that the advantage of concealment at this season as practiced by other crustacea doubtless applies here as well. The entire shell was covered by a slimy soft mucus and its colors were brighter than before, the under side of the legs showing also more blue. As seen from be- low, the flesh of the abdomen had a peculiar coagulated appearance. The new carapace had a length of 57 mm. and the old of 43 mm., and had enlarged in diameter from 25 to 28 mm. The east shell had one broken antenna, 53 mm. long, which was replaced in the new shell by a perfect antenna 68 mm. long. A week later this crayfish was active and keenly seized and ate a large Iumbriculus. Two more males cast their shells June 15 and one of them was attacked by others when only the carapace had been shed and the abdomen was as yet in its old shell; one-half of the thorax and part of the abdomen were de- voured. The other large crayfish was able to flap its abdomen vigorously when lifted out of the water though its body anid chele were soft and flaccid. In both east shell and new one the rostrum was broken off and had evidently not been regenerated. Still another large male, received in October, 1904, cast its shell May 19, 1905, and could move about though still soft. The breeding season of these crayfish was far advanced when they were received in February. In many eases the males had no sperm left in the defer- ent ducts and the females had laid their eggs, which in four.dead and six live specimens formed large dark masses attached to the abdominal limbs, or pleo- pods, and to the sternal hairs of the abdomen. Contrary to expectations, these eges were still alive and it was found possible to rear them and to get the sub- sequent stages in the life history as described below. In the hope of getting light upon the early part of the breeding season, another lot of thirty-seven crayfish of the same species were got from the same place October 29, 1904; but here again the beginning of the breeding season had passed. The only two sur- vivors on arrival were both males, and three others recently dead were also males, so that here again the vitality of the males seems to exceed that of the females. In all there were twenty-two males and fifteen females. Five of the females had eggs upon the pleopods and these were in early cleavage, showing some twenty nuclei migrating to the surface. It would thus seem that egg-lay- ing takes place in the autumn, probably in October, and subsequent observations 12 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS showed that the eggs are carried by the female all the winter and hatch in the spring. The eggs on each female were about 500 in number and formed a dark brown or more usually nearly black mass all over the under side of the abdo- men. Hach egg was very large, about 2.5 mm. in diameter, and as in other kinds of crayfish, enclosed in a complete capsule of hardened secretions that ex- tended as a slender stalk to fix each to the seta on the pleopods, or, in some cases, to the set on the sternal ridges of the abdomen. All the eggs seemed in good condition except a cluster of four or five among the brown eggs of one female and these few were overgrown with a fungus. The darker eggs plainly showed embryos in the stage represented by Reichenbach (’86) as J, fig. 12, standing out as a whitish area on one side of each egg. Thus the embryo had already advanced to a condition in which the embryonic area occupied a considerable part of one-half of the spherical egg. There was still a wide margin between the appendages and the well elevated wall that surrounded all the posterior part of the embryo. The eyes, two pair of antenna, mandibles, maxille, maxillipeds, and five periopods were well inarked. The posterior four periopods and the abdomen projected forward over the thorax so that the abdomen reached to the first maxilla. The eggs of one female showed in addition to the above embryo many nuclei seattered over the nonembryonic areas of the egg and plainly seen against the brown background of yolk. In one female the eggs were covered by a dark deposit that had to be o5 scraped off before the glossy egg capsule and the contained embryo could be seen. A few eggs were greenish and covered by a deposit that could be scraped off; when these were opened, or when boiled, the contained embryo was found to be in the stage H of Reichenbach. In one black egg the heart of the embryo was seen to beat very faintly, and after the eggs had been kept in water twenty-four hours many eggs showed the heart beating and were kept in the hope that they would develop. In water about 9° C. the eggs that were still attached to the abdomen of living females did develop, though very slowly, as will be seen from the following results. After nine days, March 2, the embryos were perceptibly enlarged with longer antenne and abdomen, the second antenne reaching back nearly half way to the end of the limb-bearing region. The heart, now lying in a plane at right angles to the ventral surface and above the base of the abdomen, was beating so strongly as to give a decided jerk to the thoracic limbs. In one embryo it beat at the rate of 66 to the minute and in another at 82. Inside the outer egg capsule there was evidently a delicate inner membrane investing the embryo. For six days more the only change noted was a slight increase in size and the extension of the second antenne beyond the middle of the limb-bearing region. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 13 Four days later a marked widening of the anterior part of the embryonic area had taken place and the slow growth of the abdomen had brought its tip up to the posterior edge of the mouth. In another week the still larger em- bryos had antennz reaching nearly to the end of the depressed, limb-bearing, region and the abdomen reached forward over the mouth as far as the bases of the second antenne. Embryos of this age, some twenty-five days after they were received, were very attractive objects seen through the transparent egg capsules. The trans- parent limbs stood out strongly contrasted against the dark red yolk mass that still took up the major part of the spherical egg. On each side of the egg the boundary of the yolk mass was slightly incised where the ‘‘liver’’? was forming. When such embryos were plunged a moment into boiling water and then put into cold water a mere seratch of a needle sufficed to cause the tough outer capsule to open with explosive force and the embryo was readily removed, leav- ing even its thin membrane sticking to the inside of the capsule. During another week there was but little increase in the length of the ap- pendages, but through the transparent walls of the limbs and body a few blood corpuscles were seen passing along in the Jarge median thoracic artery and in the antenne and periopods. Ten days later, April 5, when the water had risen from 9° to 11°, the size had markedly increased and the embryo instead of being restricted to a flat region upon one side of the sphere could no longer be seen from a single point of view, since it now extended in a curved surface over one-half of the egg. The long antennze now met one another and their tips overlapped at the deep notch where the abdomen joined the thorax. The legs and chele had grown long enough to overlie the abdomen and to conceal its tip. Such embryos were nearly in the stage K of Reichenbach and were prettily colored. The carapace had bright red pigment along its ventral border, the dark red-brown yolk took up less than half the bulk of the egg and was divided by a deep fissure into an- terior and posterior lobes which were encroached upon by the ‘‘liver’’? which was conspicuously colored. In one its contents were red and in fifteen green- ish and translucent or else white-yellow and opaque. The large eyes had also some pigment formed in them. Eleven days later the elongated walking legs and chelw reached forward over the posterior edges of the eyes. Pigment cells were as yet not seen in the above limbs, but were conspicuous in the first and second antenne and in the abdomen as well as the thorax. The delicate inner membrane was seen loosely investing the tips of the chele like a cast-off exoskeleton. Some six days later the embryo had passed beyond the stage K and was nearly ready to hatch. The limbs were even longer, so that the chela reached over part of the eyes and the antenne overlapped one another the whole width 14 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS of the abdomen in such a way that the left passing posterior to the right had its tip on the right side and the right passing anterior to the left had its tip on the left side. While all the eggs were still alive and had been kept well aérated by the swaying movement of the pleopods that the mother makes, seem- ingly for this purpose, yet some of them showed the effects of such a very long existence in the water by being covered over upon the outside of the egg cap- sule with deposits which in some cases were mixed with growths of minute fungi and in some with vorticelle. The eggs seemed under tension and burst open when thrown into hot Woreester’s liquid; the touch of a needle to the live ege also caused its capsule to pop open. As seen under a pocket lens, these eggs nearly ready to hatch present a most attractive appearance. The carapace is spangled with branching, ver- milion, pigment cells that are especially numerous along its ventral edges and near the base of the abdomen, which is broad and also well pigmented with the same kind of cell. The antenne and antennules have both red and blue pigment cells on their basal part, but these cells are not numerous and are entirely absent from the filaments. The chelw and walking legs have some few scattered pigment cells. The still large dark red-brown yolk mass has the forked, light-colored, ‘‘liver’’ projecting into it. The eyes are darkly pig- mented in all their central parts, while the surface is still clear and transparent for some distance inward. The actual hatching of the eggs took place on one female April 25-27, and on another May 1-6, and was prolonged over several days, that is, not all the eggs on one female hatched at the same time, but whether this is normal or in- duced by the artificial conditions remains to be found out from study of these crayfish in their native waters. These eggs had thus required 62 to 64 days in one case and 67 to 71 days in the other to develop from the well advanced embryo of stages H and J of Reichenbach to the hatching larva. The tem- perature of the water had slowly risen from 9° C. to 14° C. When a female died before the eggs hatehed, it was found possible to hatch the eggs by cutting off the pleopods of the mother and fastening them to pieces of floating cork so that the eggs would be suspended in well aérated water. In hatching, the egg capsule burst open over the back of the embryo, and usually opposite to the egg stalk, and then the embryo slowly glided out back- ward, much in the same way as has been described for Cambarus (Andrews, 04). In all crayfish, and in many other crustacea, the eggs remain firmly fastened to the mother during the whole period of development and when the embryo escapes from the egg-shell the old shell remains still fastened by a strong stalk that is stuck to the maternal sete. In this Astacus as the em- THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 15 bryo slowly emerged from the egg capsule it was evidently in a very inactive, helpless state, soft and unable to use its limbs, so that one might expect it to drop away from the ege-shell, fall to the ground, and continue its life, if at all, away from the mother. However, it is well known that young crayfish remain for some time upon the abdomen of the mother, which they do not leave till they are well able to swim and to walk. This period of interrelation between mother and offspring deserves special study and we will deseribe more in detail than has hitherto been done for any erayfish the remarkable structures used in en- suring the connection of mother and young from the moment of hatching up to the time of real independence and free life. Instead of dropping away entirely from the egg-case, each soft, helpless, larva hung attached to the inside of the egg-case by a delicate thread which was firmly fastened at one end to the inside of the egg-case and at the other end to the telson of the larva. The soft, pink-colored larve thus at first hung out from the egg capsules like the pulps from burst grape-skins and were then pre- vented from entirely falling away from their capsules by these threads, so that they suggested the seeds of the ‘‘cucumber tree’’ dangling out of their -pods. As the young remained limp and helpless for some time these ‘‘telson threads,’’ as we mmay call them, appeared to be of great use, since without them the larve would have fallen to the bottom of the water and having lost connection with the parent have had small chance of survival, lacking the protection or aération furnished by the mother. Such telson threads are doubtless found in the European Astacus, for in a footnote added by M. Robin to Chantran’s paper (’70), we read: ‘‘J’ai pu constater, 4 l’aide du microscope, comme |’a montré M. Chantran a |’Aecad- emie, que les petits restent pendus sous |’abdomen de la mere, par |’interme- daire d’un filament hyalin, chitineux, qui s’etend d’un point de la face interne de la coque de l’ceuf jusq’aux quartre filaments les plus interns de chacun des lobes de la lame membraneuse médiane de l’appendice caudal. Ce filament ex- iste déja lorsque les embryons n’ont encore attaint que les trois quarts environ de leur developpement avant l’eclosion.’’ And the same general fact is men- tioned in the report of the committee awarding the Montyon prize to Chantran (C. R., 75, 1872, p. 1341). Of the above passage Huxley (’80, p. 352) says: ‘‘Is this a larval coat? Rathke does not mention it and I have seen nothing of it in those recently hatched young which I have had the opportunity of examin- ing.’’ The exact mode of attachment of this filament, or telson thread, and its probable nature will be described below in connection with the telson of the larva of this American Astacus and later on in considering the like structure in Cambarus. 2 The size of the larva in its first stage is indicated by the following rough 16 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS measurements of preserved specimens: Length of head-thorax, 4 mm.; width, 3 mm.; depth, 3 mm. Length of abdomen, 5 mm. Length of antenna, 6 mm. Length of chela, 5 mm. Returning to the account of the hatching young, it is to be noted that the activities of the young were but slowly acquired; here and there amongst the mass of eggs and young upon the female some larve showed feeble movements of the scaphognathites and, later, rhythmie respiratory movements; the long antenne projecting into the water moved somewhat, the legs and chele some- times moved and the claws opened and shut. Upon escaping from the spherical egg-case the larva became but little straightened out and remained essentially a spheroidal head-thorax with a weak abdomen bent in under it and with soft, pendent limbs. However, in from one to six hours the limbs reaching about, the claws opening and shutting and the abdomen sometimes flapping up and down, it was seen that the chele managed to get hold of the stalk of the ege- case. Henceforth the larve held fast by the chele though for a time still fastened by the telson thread also. The pleopods of the parent were now covered over with a mass of flesh- colored young, showing slight movements and conspicuously marked by the two-lobed, red yolk mass, by the dark eyes and by the yellowish ‘‘liver’’ areas. The dark yolk masses showing through the pale bodies gave somewhat the gen- eral appearance represented in figure 2, which was made from a photograph of a living female shortly after the eggs (excepting one) had hatched. When forcibly torn loose from the mother the recently hatched larve, too spheroidal to rest on their ventral side and unable to stand on their legs, lay for two days on their sides, kicking their legs but unable to walk; when, how- ever, much disturbed, they managed to swim forward along the bottom of the dish by flapping their abdomens, though they still remained on their sides. When offered a piece of rough string, such young seized it and remained sus- pended in the water, holding fast by their chelw. In this way some larve were kept suspended in running water and successfully carried into later larval stages away from association with the mother. This tending to seize hold with the chelw is accompanied by a tendeney to push far in amongst the general mass of young attached to the pleopods, so that in a few days all the young are densely crowded together in a compact mass and their long chele are seen to reach far in and to be fastened either to the stalks of egg-cases or to the coagulum that binds the sete together on the pleopods. Generally both chele grasped the same egg stalk but not always and one larva was seen holding by one chele to an egg stalk on one pleopod and by the other chele to an egg stalk upon the next pleapod. As the rhyth- mic movements of the pleopods continued after the young were hatched, this larva was in danger of having its chele stretched apart. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS Wa The fixation of the chele was a gradual process; at first the claws of the chele were not opened even when those of the walking legs were opened and shut, but soon the chele claws opened and shut and soon seized hold of any solid object accessible. Sometimes the object seized was again let loose but be- fore long the chele had reached in among the mass of young and eges far enough to find and fix upon one of the egg stalks, which are of a material and size seemingly well fitted for the attachment of the claws. Henceforth the chele seemed to remain always fast and their structure as described below indicates that onee imbedded in the material of the ege stalk the tips of the claws could seareely be liberated by the crayfish. This fixation was first made out in the English erayfish by Huxley (’80), who inferred that when once fast it would be difficult, or impossible, for the claws to open again. This use of the chele to obtain attachment to the mother was exercised with what appeared to be very earnest effort and once successful it was found that the telson thread was soon ruptured. And then if the larvae were dis- turbed they flapped their abdomens up and down and it was seen that the tel- son thread had broken so that a piece of it was still attached to the telson and was waved: about by the telson like a bit of rag fastened to it. Henceforth the young held fast by the chelx only. The general appearance of this mass of young on the mother was peculiar since the rounded head-thorax was the chief part visible in each and this was of light color with a striking bilobed or horseshoe-shaped, dark red, yolk mass across it. The legs and abdomen were concealed and the dark eyes were gen- erally out of view. With bent heads and outstretched limbs their attitude ludicrously suggested one of supplication. Thus they remained for some days. When disturbed the young made tramp- ing movements with their legs but did not move from the place to which they were fixed by their long chele. The abdomen, carried down under the thorax somewhat as in the embryo, was not readily moved but with sufficient stimulus from a needle point was flapped rapidly back and forth. When a larva was foreibly pulled off from the mass its chelw, still attached, were stretched out to their fullest extent and when the larva was released the chele contracted and made the larva spring back into place where its limbs and abdomen were again drawn in under the thorax and the creature beeame again one of the herd of ‘bison’? presenting only their humped backs to the observer. If by stronger pulling the larve were torn loose from the mother the chele parted from the egg stalks without breaking and reaching about seized hold of adjacent objects such as the antenne of other larve. Left to themselves on the mass, these separated larve soon got back again amongst the crowd; but if put upon the bottom of the dish they did not yet stand up but only gyrated about by flap- ping their abdomens. 18 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS The long antenne remained for the most part low down amongst the gen- eral mass and did not yet project up above the level of the rounded backs; they were, however, carried out in front and not, as in Cambarus affinis, tucked in between the legs. Fastened thus to the mother, the larvae remained some days and then cast- ing off their shells passed into a seeond larval stage which also lived upon the mother. In one female kept in running water at 17° C., the duration of the first larval stage was only four days, but some young of another female which were kept suspended from strings hanging in water in a warm room remained five to seven days before changing to the second stage and their fellows kept in water at 14° C. remained eleven to thirteen days in the first stage. Dur- ing this long existence in the first larval stage the only change noted was a slight darkening of the color which, owing to the scattering of bright red, branching, pigment cells on a white background appeared to the naked eye flesh-color. 3efore considering the transition from the first to the second stages we will deseribe the details of the external form and appendages of the first larva. A side view of the living larva (fig. 3), suggests embryonic incompleteness in that the antenne, abdomen, and limbs are carried downward in a way not adapted to locomotion, while the globose cephalothorax and large eyes with short stalks are features of an embryo rather than of an active larva. The dark mass seen in the figure was the still conspicuous red yolk mass which from the dorsal view (fig. 4), was balanced right and left in the anterior half of the cephalo- thorax. In life the larva was translucent yet brilliantly colored by the seat- tered pigment cells indicated in black in the figures and which were absent only from the terminal filaments of the antenne and from most of the segments of the legs. In the side view the first and second antenne are conspicuous, the three maxillipeds are seen in part, the chele are very long and heavy and the four walking legs are long and weak. The abdomen bears only four pairs of pleopods and these are small, weak, and bifid. The first and sixth pleopods are not seen and the abdomen ends in a simple telson in place of the locomotor fan of the later larvee and adults. The larva is evidently very defective in locomotor apparatus, has its sensory organs not perfected, and is specialized in its strong clinging organs, the chele, and in its large digestive apparatus for utilization of the stored-up yolk. It is still essentially embryo-like in structure and in mode of dependent life, but is exposed free to the water. The same general features are shown in the dorsal view (fig. 4); which shows the split-open egg capsule and its stalk, connected by a slender thread to the telson of the larva, a ‘‘telson-thread’’ that is fast at one end to the peculiar fan-like telson of the larva and at the other end to the inside of the ruptured egg capsule. It will be noted that the head-thorax though globoidal is considerably elongated and does not have the swollen sides shown in Huxley’s THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 19 figures of the English Astacus, which, however, were doubtless drawn from aleoholie specimens, and in A leniusculus the action of alcohol is to cause great distortion of the branchiostegites. As compared with the adult, however, the proportions of the head-thorax are embryonic and there must needs be much greater elongation as well as lateral and vertical changes to bring about adult proportions. While both from side and top views (figs. 3 and 4), the characteristic rostral spine of such crustacea seems absent, full front and diagonal side views (fig. 5) show the rostrum to be well developed and armed with lateral spines, but so bent down between the eyes as to be of no such use as a de- fense as it later will be in active stages of the larva. The habit of the first stage which clings to the parent is thus correlated with imperfections of defen- sive armament as well as with presence of food yolk and imperfections of loco- motor organs. Among the latter characters may be reckoned the smooth sur- faces of the limbs and absence of sete that later will increase the areas of resistance for striking against the water as well as furnish means of sense perception. The lack of sete represented in figures 3 and 4 is still more strik- ing in enlarged views of the limbs and is in strong contrast to the hirsute char- acter of all parts of the active larve and of the adults, and this absence of sete gives the larve a decidedly embryonic appearance. The pronounced incompleteness of locomotor organs is also associated with the shortness of the thoracic region bearing legs; thus the chelw arise farther from the anterior than from the posterior ends of the head-thorax and leave little space for the walking legs, while the anterior region containing the yolk is greatly developed in size. Next taking up in sequence the nineteen pairs of appendages of the adult we find them represented in the first larva by seventeen pairs that have in the gross, as made out by Rathke for embryos about to hatch, the essential mor- phology of the adult appendages but lack the sete and differ in proportion as will be seen from the following account and illustrations. The whole exterior of the larva in its first stage is covered by a chitinous exoskeleton of such resistance that when the young were thrown into Wor- cester’s liquid they did not die for several minutes; the appendages were cut off separately from such hardened embryos and gave the views represented in the following illustrations. The first antenna stands out horizontally in front of the head (figs. 3, 4) and is straight; as seen in figure 6, it has three basal segments, five in exopo- dite and in endopodite. The segmentation of the endopodite is very obscure. The terminal segment in both endopodite and exopodite bears three obscurely pointed spines, one of which, in the exopodite, is long and apparently of the same character as the sensory sete found there in later stages. On the long 20) THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS basal and next segment of the protopodite there are a few blunt spines as in- dicated in figure 6. This also shows the auditory organ as an open pit on the upper surface of the basal segment, in the part that is swollen out laterally. The second antenna (fig. 7), though it has a long filament, is still embryonic in proportions. The base consists of two short segments, the first of which bears the very large prominence that faces inward and has the opening of the green gland, or kidney, within its depressed top. The second segment bears a few blunt distal spines of large size. The exopodite is a very large flat seale ending in a blunt point and bearing some sixteen blunt spines along its serrated inner and anterior edges. The endopodite consists of three large basal segments and of a long round filament of many segments, forty-five to fifty, the first of which is long and slender, while the followimg ones are each about one-third as long as the first. The terminal segments are again more long and slender. As indicated in figure 7, there are a few spines at the distal edges of some of the terminal segments and of some of the others near the tip. In the adult there may be 125 segments. The mandibles (fig. 8) have a well developed eutting edge which, however, is smooth and not toothed as it is in all later, functional, stages and there was no evidence found that these organs were used. The palpus has three segments and is smooth except for the distal face of the third segment which is sparsely set with rather acute spines, many of which are shown in figure 8, and more of which are present upon the inner aspect and not seen from this point of view. The palp is thus a blunt club with terminal spines. The first maxilla (fig. 9) is very small and made of two flat plates and a somewhat rounded and blunt endopodite of curved finger-shape. Here for the first time we meet with a few small, plumose setx along the outer edge of the distal segment. The two flat plates that represent the protopodite are spinulous at the ends, much as in the palp of the mandible. The distal piece is also armed with a row of a few spines along its proximal edge as seen in figure 9. The ends of both plates are set with spines on the face toward the mouth; the proximal plate is rounded, the distal one truneated to form a jaw-like organ. The second maxilla (fig. 10) bears the long scaphognathite which has a dense row of plumose setz all along its extensive free edges. There are also a few plumose set at the base of the endopodite, as in the first maxilla (fig. 9). As above noted one of the first muscular activities aequired after hatching is the slowly developed rhythmic beat of the seaphognathite, and with this use of this appendage there is present an armament of plumose sete lacking else- where in the locomotor organs of the larva. While these plumose sete are not used in locomotion their function as flexible areas of resistance to the water which the scaphognathite bales out of the branchial chamber is akin to that of locomotor sete. The rest of the second maxilla is nearly bare of sete, but there THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 21 are a few, long hair-like sete standing out from the external face of the pro- topodite. The protopodite has the same two plates seen in the first maxilla (fig. 9), but each is deeply cleft, so that four free tips project toward the mouth. The four tips each bear blunt spines on the outer and distal faces as shown in figure 10, while upon the inner faces toward the mouth they are all set with longer, sharp, curved spines that are claw-like. The endopodite though longer and more slender than in the firstmaxilla is still very simple and not seg- mented. The first maxilliped (fig. 11) shows the protopodite again reduced to two flat plates much as in the first maxilla (fig. 9), and there are simple spinules on their cutting edges. The endopodite is small and simple, intermediate be- tween that of the second and first maxille. In place of the seaphognathite there are two movable parts; a long flat plate, the epipodite, which is without sete though sparsely spinulous on its posterior edge; and a very long and prominent exopodite. The exopodite has a very long swollen basal part with very long plumose sete on its outer edge as shown in figure 11 and is else- where naked. The terminal part is long, slender and with a very few spines at its tip. As the base of the exopodite lies over the distal end of the epipodite it is not readily seen that the epipodite has a short truncated extension sug- gesting the anterior end of the scaphognathite as well as the evident posterior blade that reached back into the gill chamber and is comparable to the like portion of the scaphognathite. The second maxilla (fig. 12) is more complex; the two segments of the pro- topodite are subordinate in mass and extent to the greatly developed en- dopodite and gill structures, but they bear a few plumose sete upon their inner edges. The endopodite resembles that of the first maxilla in position, general form and curvature, but is not only larger but subdivided into five seg- ments and bears spines. In addition to the spines shown in figure 12, there are also long curved ones on the inner face of the terminal segment. Compared with that of the first maxilliped, the exopodite of the second has a very narrow basal part which is without plumose sete but bears a few long spines on its external edge. The epipodite is present as a long, curved lamina, bilobed at the tip, and along its inner face are borne the numerous blunt filaments of the gill, podobranech. This podobranch is free at its tip, but elsewhere adnate to the lamina and bearing two rows of blunt side papille or gill filaments which are directed toward the apex, and increase in size in each row from base to apex. The epipodite lamina has a few plumose setx on the rounded ridge at its base and along its edges are scattered curved, short hooks, while its emarginate tip bears a few blunt, fringe-like papille. In addition to the above gill there is one arthrobranch that is shown in figure 12 to have a long slender stem ending bluntly and bearing two rows of blunt, curved, finger-like lateral filaments 22 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS which may each end with a small, blunt spine. The third maxilliped (fig. 13) exhibits the typical morphology of this organ in the adult; a somewhat two jointed basal protopodite bears a large five-jointed endopodite of great size, a long slender exopodite, a large epipodite and podobranch; and two arthro- branchie arise from the region connecting the appendage to the body. The endopodite bears spines upon all its segments and the protopodite has a couple of small spines upon its distal segment and a plumose sete upon its proximal segment. The exopodite, in contrast to that of the second maxilliped, has a shorter and more slender basal segment devoid of spines, while the second segment has several spines at and near its tip. The lamina of the epipodite has the same characters as in the second maxilliped, but the plumose sete along its basal ridge are twice as many. The podobranch is like that of the second maxilliped. Of the two arthrobranchie the anterior one is much like its homologue on the second maxilliped, while the posterior one, nearer the observer in figure 13, is smaller and more simple with fewer lateral filaments. Coming next to the ambulatory appendages, we find the usual large chele, the two pairs of slender chelate and two pairs of non-chelate legs (figs. 14, 15, 16, 17, 18). In these appendages of the first larva there are, as in the adults, no exopodites, and even the remarkable exopodite sete of later larve and of adults are absent in this first stage, thus adding to the simplicity of the limbs, which is also expressed in the entire absence of plumose sete and the presence of but few sharp spines. The chela (fig. 14) has the recurved terminal hooks first made out by Hux- ley in the English Astacus, and which lead to the firm locking of the chele to the egg stalk, as above narrated; and the opposing edges of the claw are ser- rated from the presence of sharp spines pointing toward its tip. The chela bears very large sharp protuberances along the inner edge of the meropodite segment, of no apparent use, while the great length and thickness of the whole limb is apparently necessary in that firm holding of the larva to the mother which resists the force of the maternal pleopods that swing the larve back and forth. The epipodite and gills of the chele are like those of the third maxilliped. The following two legs (figs. 15, 16) are like one another in every way except in proportion, the first being shorter than the second. Fach has a sharp claw with spines pointing toward the tip, but the tips are not recurved as is the case in the big chele. The gills on these two appendages are like those of the chela, but there is in addition a slender simple gill upon the body wall near the arthrobranchie. This pleurobranchia is a single filament with no lateral outgrowths and may be regarded as rudimentary at this stage, as it is also in the adult. Phe remaining legs (figs. 17, 18) have terminal segments almost like those THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 23 of the preceding legs, but there being no opposing outgrowth of the propodite, there is no claw. The legs increase in length and in slenderness from before back in the series behind the chelw. The penultimate leg (fig. 17) has a longer pleurobranch associated with it but otherwise its gills are as in the preceding limb. The last leg, however (fig. 18), has its gills suddenly reduced; in place of the epipodite and podobranch there are but a few plumose hairs such as stand upon the basal ridge of the epipodite of the preceding somites. The arthro- branchs are entirely absent and there is but one gill which is a pleurobranch, which, however, in place of being a simple filament or rudiment, resembles a re- duced or simple arthrobranch in that it has about seven short lateral processes in two imperfect rows. The branchial formula of the first larval stage of Astacus leniusculus is then as is given in the table below. This was found to be just the same in the adult of this species and it is said to be the same in the English Astacus pal- lipes, except that the latter Jacks the rudimentary pleurobranch on the somite of the first leg. Arthrobranchie. | Podo- Pleuro- | ip tal branchie. | | branchiee. | eee Anterior. | Posterior | =a | — a ae eee Somite of 2d maxilliped 1 | 1 | 0 0 | 2 Somite of dd maxilliped . . . . .| I 1 1 0 3 Somiterofichelay =. 2 = 5 = . . «| 1 1 1 0 3 Somite of Ist leg 1 1 1 1R 8+1R Somite of 2d leg 1 1 | IR 3+1R Somite of 8dleg . . eee? i 1 1 | | IR 3+1R momiterotAéthylesss 5 5 5 2 « ss 0 0 0) | 1 1 6 6 5 IeEo RN 1esnsiR The above illustrations of the separate appendages of the head-thorax are the first ones as yet given of any larval Astacus, since the previous illustra- tions of European forms are only the small mawxille and maxilliped of an em- brvo not vet hatched as depicted by Rathke (’29, fig. 29), the tip of the chela shown by Huxley (’80, fig. 8), and the under side of the abdomen with its pleopods figured by Reichenbach (’86). Upon comparing the adult appendages of Astacus leniusculus with those of first larva as above described, the fundamental agreement in morphology was obvious, but there were the following differences which all suggest a linger- ing on of embryonic characters into the life outside the egg-shell. Throughout all the appendages there was a marked lack of sete correlated with evident lack of locomotion and probable weakness of sensory activity. Excepting the chele the cephalothoracic appendages had no obvious use. The first antenna, having 24 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS but five segments in its exopodite and in its endopodite, is but embryonic com- pared with the adult that has at least thirty-five in its exopodite and almost as many in its endopodite, a difference which is of great moment when we con- sider the repetition of special sense organs that are found on many successive seements of the exopodite of the adult. The auditory organ also is apparently of no functional value in the early larva.* The number of segments in the fila- ment of the second antenna must also increase greatly to’ form the one hun- dred and twenty-five of the adult; an increase apparently brought about by in- terpolating new segments at various places by division of the old ones into two. While all the adult appendages of the head-thorax are represented in the larva at hatching, this is not the case for the abdomen, for the sixth pair of ab- dominal appendages are not externally present and the first pair which in the adult male are so essential are absent in the first larva as they also are in the adult female. The other abdominal appendages are four pairs of simple ple- opods which hang down beyond the pleural plates of the abdomen so that they are seen from a side view (fig. 8). Each pleopod (fig. 19) is as in the adult composed of a short and a long segment that make up the protopodite and of two simple terminal plates, the endopodite and the exopodite. These are slightly curved and armed at the ends and to some extent on the edges with small weak spines and they entirely lack the plumose sete that makes them use- ful in the adult for fanning the water. The illustration is of the anterior face of the left pleopod of the second abdominal segment and shows that the ex- opodite is longer and wider than the endopodite, while in the adult the exopodite is much the shorter and smaller in the female pleopods that bear the eggs and in the male pleopods that transfer sperm. The appendages of the sixth somite though not externally free are yet present and of large size though imperfect in development and lie within the substance of the telson, as can be seen in transparent living larve. It is their presence which swells out this region ventrally and gives rise to the protuberance seen from the side view (fig. 3). Looking at the telson from above (fig. 20), the very imperfect future sixth ple- opods are seen as two somewhat less translucent areas, right and left in the anterior part of the telson and each having an outline suggesting that of a mit- ten. Only later will these concealed buds of the sixth appendages burst out after a moulting and expand as the very large lateral parts of the caudal fan, so essential in quick locomotion, This retention of these appendages within the telson in Astacus was known to Huxley (’80), and figured by Reichenbach (’86). The telson requires special consideration in connection with the ‘‘telson- thread,’’ as mentioned above. The telson itself is a very large plate with nearly circular outline and is thin posterior to the above region occupied by THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 29 the buds of the pleopods on either side of the intestine and anus. In structure it is essentially a translucent, vascular, connective tissue mass over which are scattered brilliant vermilion pigment cells, indicated in black in figure 20, and which is covered by a thin epidermis and thin chitinous cuticle. Along most of the free margin of the telson is a row of blunt, stiff papille, or spines. in all there are about sixty-six of these spines arranged symmetrically, half on each side of the median plane. In the figure of Astacus given by Reichenbach (’86) there are but twenty-five spines or dentations on each side of the median line and this may well be a character of systematic value. In addition to the thirty-three lateral spines of Astacus leniusculus there were six or seven smaller, blunt spines on each side which stand in between some of the larger ones, one small one between two larger ones, and generally not so near the edge but more up on the dorsal surface of the telson. The interior of the telson has a radiated appearance like that figured by Remak and by Reichenbach and which was referred by Huxley to the disposi- tion of vascular canals; but in our present larve this radiation is due to long delicate lines passing centrally inward, one from each lateral spine and from some of the smaller spines, thus making the divergent system radiating from near the anal region as shown in figure 20. Subsequent events show that these lines are the forming plumose set for the perfect locomotor telson of later larval stages. Hach when enlarged (figs. 21, 22) is a bundle of fibrille that are very small in comparison with the nuclei of the epidermis as each bundle of very many fibrils is but once or twice the diameter of a nucleus. These radiating lines are in fact to be likened to compressed bottle brushes which later will expand as the perfect locomotor plumes on the telson of the second larval stage (fig. 23). The plumes are being made within epidermal tubes or glands and between the successive radiating glands the vascular spaces, also radiating, form the justification for Huxley’s interpretation. During the first hours of larval life the telson is connected to the inside of the egg case by the long telson thread represented in figure 4. This is a trans- lueent, chitin-like membrane, or flat ribbon, showing a striated appearance due to fine wrinkles in it (fig. 20), but otherwise apparently homogeneous. Though seemingly but superficially attached to the telson the contact is a very firm one so that when the larva succeeds in getting hold of the egg stalk and finally flaps its abdomen strongly enough the telson thread is broken before it is torn loose from the telson. When enlarged (fig. 21) the mode of attachment of the very thin but tough membrance is seen to be, that ten of the marginal spines of the telson bear special hook-like projections that are fast to the membrane. These few spines are different from the others though some of the adjacent ones have somewhat of the same structure at their tips. While in figure 21 the spines and their processes are represented in black, they are in nature trans- o oO 26 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS parent, colorless and very inconspicuous, especially the processes which are hyalin, myelin-like protrusions, suggesting the products of glandular activity. While many of these protrusions end bluntly others seem to be contimued as fine thteads that are fast to the membrane or telson thread. Moreover, many of the protrusions bridge over the space from one spine to the next and are con- tinuous with adjacent protrusions as if they had flowed out when viscid and then coagulated. Such bridges make adjacent spines into hooks that hold fast to the membrane, but the ultimate and essential fastening of the membrane to the telson is by fine threads of some coagulated material that seems to be a con- tinuation of the grosser protrusions figured in black in figure 21. It would appear from the statement of M. Robin cited above (page 15) that in the Astaenus studied by Chantran there are but eight spines used for attachment to the telson thread. The origin of these peculiar glandular spines, for such they seem to be, is to be sought in the embryo. At the time of hatching the egg capsule breaks and for a brief period the embryo may still be enveloped in a very delicate membrane which passes over the abdomen and all about the telson. At that period the spines of the terminal part of the telson abut against the invest- ing membrane as shown in figure 22, which is from a specimen just hateh- ing and killed in Pereny’s liquor. Here are shown the epidermal nuclei, the striations, or forming plumes of the future telson, continuing up through the body of the spines, only three of which are drawn; and at the tips of the spines fountain-lke masses of blunt protrusions, swollen at the tip and in many cases pressed against the membrane. It would seem that a viscid mass had been poured out from the tips of the spines and that this oozing out in threads had become firmly soldered to the membrane. As far as made out the origin-of the supporting telson thread is thus as follows. The thread is really a membrane and when the embryo is hatching this membrane is spread like a loose skin all over the embryo inside of the egg case. When the embryo hatches it also sheds this membrane, coming finally to pull its abdomen out of the part of the membrane that surrounds the abdomen like a long bag. The bottom of the bag is, however, fastened to the tip of the telson as indicated in figure 22, so that the creature cannot get entirely free from the bag but pulling out its abdomen pulls up the bottom of the bag and turns the bag inside out. The struggles of the larva drag the membrane into a long thread, and one end of this remains attached to the tel- son spines as seen in figure 21. Once this is accomplished the fact that the clear thread is really a cast-off membrane would not be suspected, since it seems a homogeneous, finely wrinkled thread that might well be a secretion. The similar structure in the European Astacus was mentioned merely as a filament as cited above, page 15, but Huxley intuitively queried if it might be a larval skin. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS Dit The other end of the telson thread remained fast inside the egg capsule (fig. 4), and this attachment is as important as the above described attach- ment to the telson in making the thread of use to secure the larva from being lost. How the connection of the inner membrane to the egg capsule was brought about was not determined but it was existent long before the embryo hatched. Embryos three days before hatching killed in Worcester’s liquid and soaked a week in five per cent potash showed an outer egg capsule, an inner shell and a membrane that was loose and visible over the chelx and over the deeply bifid telson which already bore terminal spines. And embryos nine days be- fore hatching showed the same double shell and membrane, but no telson spines. The telson thread is thus a thin membrane formed about the embryo and early fastened by radiating fibrils (fig. 4) to the inside of the inner of the two layers of the egg capsule. Later in the life of the embryo this membrane be- comes also fastened to the telson by secretory activity of the terminal spines. When the embryo hatches the membrane is ruptured and in part turned inside out and drawn into a thread-like form, fastened at both ends. Other facts re- garding the telson thread will be given below in the description of Cambarus, in which it also exists. The passage from the first to the second larval stages was seen in some larve lying in the bottom of a dish, and in others that had fixed themselves to strings. In these the old larval skin burst open and the second larva, as it were, ‘‘oozed’’ out backward for several minutes and its chele and abdomen re- mained longer inside the old skin but were then suddenly withdrawn. For a few minutes the larva in its new stage lay stretched out straight, as if dead but then flopped its abdomen, moved its legs, got upright and walked and even swam backward and finally crawled up into the piles of other young in the same second stage. While the larva in the first stage was inactive and remained always . fastened to the mother, the second larva was active and finally abandoned the mother though for a time still associated with her. Upon casting off their first larval skins the larve in the second stage leave those skins fastened by their chelx to the egg stalks on the mother’s pleopods, and are free to crawl about over the pleopods of the mother amongst their numerous fellow-larve. Soon these larve descend the pleopods and make short excursions under the abdo- men of the resting mother and over various parts of the mother’s body, finally wandering off over the bottom of the aquarium for short distances to return frequently to the mother again. The mother thus had fastened to her pleopods a large mass of old egg stalks and capsules to which were fastened the cast-off skins of the larve, and over this mass erawled the active larve till after a few days the egg cases and cast skins as well as egg stalks were found to have disappeared leaving the ple- 5S 28 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS opods clean but still occupied by the larve. In the Astacus in France Chan- tran (’71) finally convinced himself that the larve ate their cast skins and the egg capsules; and the same probably occurs in Astacus leniusculus. The habits of the second larve showed much greater diversity than was possible in the attached larve in the first stage but through the early part of the second stage a tendency to climb seemed a dominant feature of their lives. When a number were put into a dish by themselves they tended to climb up onto one another to form a mass but if put back with the mother they soon climbed up onto her pleopods where they held on so firmly that when a pleopod was cut off and thrown into 70 per cent aleohol, some of the larve still retained their hold though most of them did not. When the larve alone were in a dish with a spray of myriophyllum they climbed up it and crawled together in a mass between the plant and the glass; but they did not climb up onto a piece of cotton cloth hanging down in the water from a floating cork. Even when the mother was dead, the young twenty hours after passing into the second stage continued to hold firmly to the maternal pleopods. But after three more days the young had ceased to huddle together so much, and crawling about over the bottom of the aquarium, and sometimes swimming, they were at times carried away with the current of water. Though some of the larve concealed them- selves under ooze and dead leaves at this time, others continued to hold on to the abdomen of the dead mother for four or five days, when the abdomen was cut off and fastened at the surface of running water, but about May 18 these larve also dropped to the bottom and lived there. This climbing instinct can then be satisfied in various ways, and when thirteen larve were removed and put into a dish with another female bearing young, but few minutes sufficed for five of the thirteen to find and to climb up onto the pleopods of the strange female. The possibility of resolving these habits of the young second stage crayfish into so-called tactic phenomena, into chiefly geotactie and stereotropic responses, will be considered in connection with some observations upon the young of Cambarus. The general form of the second stage as represented in figures 23 and 24 is obviously more like the adult than like the first stage as is true also of the habit. Comparing these figures with 3 and 4 there is a noteworthy change in size; after casting off the first skin the crayfish measured 11 mm. from tip of rostrum to end of telson, exclusive of the long fringe of plumose sete which made the length 12 mm. if the set were included, and so greatly enlarged the area of telson available for locomotion. The thorax was 2 mm. wide and about 2.5 mim. deep. The cephalothorax was 6 mm. and the abdomen 5 mm. long; the telson was 2 mm. wide without the set, and 4mm. with the sete. The an- tenne were 10 mm. and the chele 8 mm. long. All these measurements were taken from preserved material and show that the animal was now a large larva. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 29 Its brilliant color added to its size made it an attractive larva resembling a young lobster after the swimming stages. As indicated in black in the above figures the pigment cells were scattered over the head thorax and abdomen and more sparingly over the chelew, walking legs and basal parts of the antenne. To the naked eye the larva walking on the bottom of a dish seems light flesh-color, translucent and inconspicuous, but the chele look red, the eyes are dark and the yolk is still a very evident dark, red-black mass of bilobed form across the middle of the head thorax. The liver lobes anterior and posterior to the yolk were noticeably yellowish and greenish. The abdomen was flesh colored for the most part, but the telson was nearly colorless and with a fringe of white, clear sete so long as to suggest a peacock’s tail. On the first abdominal somite the densely crowded pigment formed a conspicuous eross-band (fig. 23). Another such aggregation of pigment was found posterior to the eye and external to the base of the rostrum. In addition to the color due to the much branched red pigment cells, indicated in black in figures 23 and 24, there soon came to be a variable amount of blue color not so readily seen and due to large blue pigment cells. In strong light the red pigment often stood forward on a back- ground of blue. The blue was evident on the basal part of the antenna and antennule, on the mandible and its palp, but not on the mavillipeds. On the dorsal side of both thorax and abdomen there were some blue, faint, scattered areas internal to the red. As shown in figures 23 and 24, the cephalothorax in passing from the first to the second stage had become long, narrow, and angular with a long gothic rostrum standing straight out in front between the eyes on a level with the back. The rostrum also had large lateral spines at its base and half way out its length. In walking about these larve carried the antenne and the red chele for- ward and the abdomen straight out behind as in figure 23, and not bent in under the thorax as in the first stage. However, when not walking the abdo- men was bent as in figure 24. As in a young lobster the slow walking was quickly replaced, at alarm, by rapid backward swimming caused by flapping the abdomen with its extensive telson fan. As the larve went about more and more away from the parent, they became more individual and more complex in their movements; they were seen to scrape the backs of their heads with their legs, to raise their chelew as if in defense when a shadow passed over them, and in other ways to act like an adult crayfish much more than did the sluggish and simple first larve. In watching one of these second larve slowly walking, the movements of the five long limbs seemed to be as follows. The fifth, fourth, and third limbs standing out at the sides of the body (fig. 23), were so bent as to hold the body high up above the bottom of the dish and swing back and forth as the 30 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS chief locomotor organs. The chele were held in readiness in front of the animal and the second limbs (fig. 23) were bent inward under the body with the tip for- ward so that in moving forward the body rode over these appendages as upon levers with very little movement of the base of these appendages. The third appendages always pointing forward swung from an angle of about 45° with the side of the body to one of about 70°. The fourth limbs had a very long swing from a forward position about 45° with the side of the body back past 90° with the side of the body. The fifth limbs had a very short, hobbled, movement like the third but always directed backward, from about 100° to 120° with the side of the body. In taking up the appendages of the second larva, in sequence (figs. 25 to 39) it will be noticed that the relative nakedness of the first larva has given place to a hirsute condition, indicated in figures 23 and 24; set occur upon the antenne, chela, legs, pleopods as well as the sides of the abdomen and, as the separate sketches show, upon all the other appendages. The active second larva has thus come into possession of sensory and locomotor setz lacking in the imperfect first stage, and similar to those of the adult stages. The first antenna (fig. 25) still has the same general form as in the first stage (fig. 6), it has five segments in exopodite and in endopodite but it is noticeably more finished in being well armed with sete. The narrow part of the proximal segment bears a sharp spine upon its inner side. The auditory pit on the basal segment is now well guarded by a row of plumose sete pass- ing from the outer edge inward and spread across the orifice of the pit. There is also a row of sparsely branched plumes along this upper face of this segment and parallel to its inner edge and in addition there are a few other set ar- ranged as in figure 25. The second and the third segments bear long plumose set on their inner sides and long, stiff spine-like sete on the external sides of their distal ends. The endopodite and the exopodite each bear a few long stiff spine-like sete at the distal ends of their five segments and in addition the char- acteristic blunt sense clubs of this appendage are now evident. These organs are placed in groups upon the inner and lower faces of the third, fourth, and fifth segments of the exopodite. The third segment has a cluster of three upon its distal edge, the fourth has a group of two at its distal edge while the fifth has a group of three at its middle part, where its diameter suddenly diminishes. As all these sensory clubs face downward they are foreshortened in the above figure and in reality are much longer than shown. The second antenna (fig. 26) has increased greatly in length over its former state (fig. 7). Its basal parts are more angular and the excretory cone on the basal segment is relatively very much smaller while the spines of the second segment are borne upon a large, seale-like protuberance. The exopodite scale bears a row of long, plumose sete all along its outer edge while in the first THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 31 stage there were only simple spines to anticipate some of these plumes. The three large segments of the exopodite are more angular than before and now bear a few sete while the filament has some fifty-four segments of the form and proportions shown in figure 26. As these bear needle-like setie on their dis- tal edges the filament seems under a low magnification somewhat like a brush. The mandible is not only greatly enlarged but more complex in having an effective cutting edge no longer smooth but serrated by seven unequal angular projections (fig. 27). The palp is more complex in having more numerous long, acicular sete both on the exterior and interior faces of its terminal segment and a very few sparsely-branched plumes on the distal part of its second seg- ment. When folded down the palp fits into a deep depression on the inner face of the mandible (fig. 28), and the proximal border of this depression is irregularly dentated with rounded protuberances. The exoskeleton over these dentations and over the sharp teeth of the cutting edge is now very thick and horny, being solid as far back as the second line in figure 28. While the acicular sete over the terminal segment of the palp appear smooth under Zeiss 2 A, they are really set with short, fine, side branches along their distal halves as seen with 2 D, and they have rather blunt points so that they would seem to aid in a brush-like use of the palp. The first maxilla (fig. 29) has progressed beyond its former stage Gigs 9); chiefly in the outgrowth of long sete in place of blunt spines and also in the addition of sete where there were no outgrowths at all. The sete are of two kinds, a few plumose and many acicular; the latter are found chiefly on the ends of the two plates of the protopodite where they replace simple spines, while the plumes are chiefly lateral. On the basal, or first segment, however, there is some- what of a transition, since its proximal border is set with sete that extend out to the tip as very long and sparsely branched plumes that thus extend close up to the acicular set. These latter under 4 D are seen to be set with very few and fine side branches so that they are really somewhat plumose. The similarly placed acicular sete upon the second segment, however, show no side branches but are really smooth. The small tuft of sete already present in the first stage at the base of the endopodite remains but little changed in the second stage. : The second maxilla has undergone like changes (fig. 30). The terminal spines of the first stage (fig. 10) are replaced by plumose set and a few more long plumes are added. Here again the plumose sete of the proximal edge of the first segment extend out as far as the acicular sete of the apex. The endopodite has a few long plumose sete on its distal part in addition to the cluster at its base. The scaphognathite is but little changed, the sete along its outer edge being so bent down that they do not show their full length in the view represented in figure 30. However, at the posterior tip of this respiratory 32 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS organ there is a peculiar growth of one or two setw. These are very long and though appearing quite smooth and hair-like, with 2 D, they are seen to be really set with short, fine side bristles, and they are sharp pointed. On the inner faces of the four lobes of the protopodite there are now many long, sharp, curved, sete. Tn the first maxilliped (fig. 31) the process of substituting plumes and acic- ular sete for spines has been carried on in the same general way. The cut- ting edges of the protopodite now bristle with sete most of which are simple, some straight, some with curved tips; but the tendency to run the plumose sete up to the tip of the basal segment is carried so far that these plumes take entire possession of this basal segment and only the second segment has acic- ular sete. Even on the second segment a few plumes come up nearly to the tip, on the distal side. The endopodite bears but a few plumose sete while the long exopodite has in addition to the former series of plumes along the outer edge of the basal part (fig. 11) several remarkably long and conspicuous plumes near its tip. The terminal part of the exopodite is cylindrical and tends to become segmented and it is from its last and its penultimate ségments that the long plumes project and form a terminal brush. The flat epipodite scale has still a few minute, blunt papille along its posterior edge (figs. 31, 11), and on its outer and inner edges a few of the remarkable exoskeletal hooks char- acteristic of these organs. The second maxilliped (fig. 32) has changed chiefly in adding plumes and sete. The gills remain as before (fig. 12) but are longer. The protopodite has added longer and more numerous plumose sete and a very few acicular ones. The exopodite has substituted for its few terminal spines a brush of several remarkably long, strong plumes (fig. 32), and for the sparse spines along the outer edge of its basal segment a few sete which proximally are sparsely plumed and distally smooth without barbs. The endopodite looks much changed owing to the development of many long and stout acicular sete over its terminal parts and inner edges. The sete of the endopodite are smooth as seen with 2 A, but with 2 D some five or six on the inner face of the second segment and again on the distal edge of the fourth segment are finely harbed. In the latter position two at the corner toward the exopodite have their fine lateral branches flattened like saw teeth, so that they resemble the cleansing sete upon the penultimate segment of the fifth leg of the adult, elsewhere described (Andrews, ’04). The fifth, or terminal segment of the endopodite, is armed with long, smooth, stout, spine-like sete, with blunt points. The third maxilliped has added but few plumes but many very long acic- ular sete (figs. 33, 13). The plumes are a noticeable bunch at the tip of the exopodite and a few on its basal part, as well as very few on the protopodite and an increased number at the base of the epipodite. The gill region is now THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS ay Ww for the first time provided with the phenomenally long hair-like coxopoditic sete found in the adult and in all but the first larval stages on this and all ‘the following thoracic somites. As seen in figure 33, these setx arise external to the exopodite and add a conspicuous element to the appendages as they are longer than the entire exopodite and coil about like stiff wires. The endopodite bristles with exceedingly long, sharp needle-like setw and has no real plumes whatever, though with 2 D it was evident that some of the longest needles on the terminal segment were very finely barbed. On the anterior, and inner, face of that segment there were also numerous stout sete with flat saw-teeth like those above described, on the corner of the penultimate segment of the endopodite of the second maxilliped. The chela (fig. 34) has greatly increased in size, as shown in comparing with fig. 14, and now has simple acicular sete seattered over it but no plumose sete. The terminal claw is no longer used as a holdfast and has no longer recurved tips. Henceforth of use as a cutting shears it now bears a large tooth on each blade as well as the rows of spines formerly present. At the base of the appendage the gills have increased in size and there is a long tuft of coxo- podite set, which, however, are short in proportion to the enormous endopo- dite. The four walking legs (figs. 35, 36, 37, 38) though much larger than in the first stage (figs. 15, 16, 17, 18) have not increased as much as have the chelxe and they retain their relative proportions and sizes. Like the chelx they now bristle with acicular sete and show no plumose sete. However, at the distal edge of the penultimate segment of the fourth and of the fifth legs (figs. 37, 38) there are a few of the saw-like ‘‘cleansing setie’’ previously referred to as oc- curring in the adult. The gills have changed only in size; the coxopodite sete are very long threads, but only few in number and on the last leg reduced to one. The branchial formula in the second stage was thus just like that above given for the first stage. On the abdomen of the second stage there were still but four pairs of ple- opods since the first were not yet formed and the sixth still remained inside of the telson. But each pleopod was now so well provided with plumose sete that the appendage simulated a locomotor organ. The larve also now had the adult habit of swinging the pleopods back and forth and so producing currents in the water which may well be of aid in respiration as they would change the water supplied to the inhalent openings of the gill chambers. Hach pleopod (fig. 39) had grown greatly in length as compared with its first appearance (fig. 19); the long plumose set arose from the distal parts of the exopodite and endopodite and resembled the plumes upon the exopodites of the maxillipeds (figs. 31, 32, 33). The exopodite was still the longer and the endopodite the 34 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS shorter of the two flat lobes that bear sete so that the adult relationship in size of these parts was not yet arrived at. While the telson of the first larva was nearly circular, in the second it was swollen laterally (fig. 23), and became thus transversely elongated since the sixth pleopods had now grown within it as very large lateral masses. The posterior edge of the telson was somewhat incised on the median plane and thus recalled the early bilobed condition found in the embryo some days be- fore hatching. The long plumose sete of the telson (fig. 23) that aid the larva in swim- ming are the expanded products of the radiating glands seen within the telson of the first stage (fig. 20). Thus provided with effective swimming sete and more numerous sensory sete the second stage larva gradually depends less and less upon its mother and finally leaves her altogether. After some eight to ten days these active larve cast off their shells and passed into a third stage. The third larva was in the main very like the second but it had advanced a very important step in freeing its sixth abdominal appendages which hence- forth are not inside the telson, but lying by the side of it to make the effective tail-fan that is used in rapid locomotion. Some hours before shedding the see- ond larva plainly showed the sixth pleopods as dark red, partly opaque masses within the base of the telson and after shedding these appendages were ex- panded as is shown in figures 40 and 42. As both the end of the telson and the edges of these great flat sixth pleopods are set with plumose sete the combined fringe of seta augments the surface used by the larva in escaping backward by vigorous blows of the telson and sixth pleopods against the water. The details of this effective and very large sixth pleopod which has been forming slowly on each side within the base of the telson ever since the larva came out of the egg, that is from some two to three weeks, are shown in figure 42, which shows the dorsal face of the left appendage of the sixth abdominal somite. This appendage joins onto the sixth somite and lies by the side of the telson as indieated in figure 40. The protopodite bears a prominent spine over the base of the endopodite; the endopodite is armed with two spines near its edge and the exopodite with five spines, along the edge of the two segments into which it is divided, as in the adult, by a movable hinge. Seattered over the sur- face are a few relatively short acicular sete. It will be noted that the plumes along the edges of both endopodite and exopodite are arranged to make a most effective fan since those of the endopodite overlap some of those of the exopodite, when, as in figure 42, the exopodite is not extended as far as possible away from the median plane. At times the exopodite may be shut in under the endop- odite like a part of a fan. The real length of the terminal plumes is somewhat greater than shown in THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 35 — the figure since they are foreshortened there; they do not stand out in straight continuation of the plane of the telson and sixth appendage but are curved downward so that their tips tend to point forward under the animal. The entire fan is thus somewhat concave on the ventral side and in life the larva carries its abdomen with its tip close to the substratum upon which it walks or stands (fig. 41), and seen from above the fan is foreshortened. Both the habit of carrying the fan inclined downward and the curved growth of the plumose sete combine to make the fan a hollow scoop, which form would seem to be a more efficient one for striking the water forwards and thus propelling the animal backward. The forward bending of the sete by making it more difficult for them to be forced back beyond the plane of the stiff parts of the fan would seem to make them a more efficient addition to the striking surface. The period of life in which the effective tail fan is formed by the lbera- tion of the sixth pleopods has hitherto remained unknown for the genus As- tacus. Huxley (’80) says: ‘‘I imagine * * * that the appendages of the sixth abdominal somite are at that time (during the first ecdysis) expanded, but nothing is definitely known at present of these changes.’? Faxon (785) records that specimen of Astacus pallipes 11 mm. long and ten days old still had the sixth pleopods enclosed within the telson and supposed that they would be set free after the second or third moult. Other observations seem to be lacking. The telson seen foreshortened in the natural position figure 40, is seen in its true proportions in figure 43. From the circular form of the first stage (fig. 20) it has passed through the transversely elongated form of the second stage (fig. 23) to its present complex and angular form. By an imperfect transverse hinge it is now divided into an angular anterior part and a rounded posterior part. The fact that this transverse division of the telson does not show till the third larval stage is of interest in connection with the fact that this seems one of the recent acquisitions of the highest crayfish. In the lobsters and other marine forms, as well as in all the crayfish of the Southern Hemisphere, the telson is not at all divided and amongst the Potamobine or higher crayfish of the North- ern Hemisphere the division of the telson is much more perfect in the highest forms, such as Cambarus affinis, less pronounced in some lower forms as Cam- barus Clarkii and in Astacus leniusculus, which is doubtless less specialized than Cambarus, the division of the telson in the adult is by no means a perfect one. In figure 43 are shown groups of acicular set symmetrically placed right and left; the rounded, terminal lobe is the only part bearing plumose sete. These plumes stand in a single row and on the right and the left begin ante- riorly as short sete followed by others that very soon are much longer and of about constant length along the posterior border till near the median line when there is a sudden falling off in length and one very short sete ends the series. 36 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS Just dorsal to this row of long plumes there is a row of much fewer and re- latively short acicular sete, shown in black in figure 43. In the reentrant angle between the two lobes of the telson, each side, there is a rounded spine similar to the one formed on the sixth pleopod (fig. 42) where the basal lobe of the exopodite joins the distal lobe. Beyond this perfection of the locomotor apparatus of the abdomen the third larva differs but shghtly from the second though there are differences in size, proportion, color and habits. Comparing figure 40 with figure 23 it is seen that the abdomen is wider, the cephalothorax more cylindrical, the limbs stouter, the entire aspect more heavy and erayfish-like. The third larva is still sufficiently transparent to allow the heart to be seen through the shell, under the microscope, beating at about three times a second; the stomach and intestines also show dimly as dark areas. The animal is not conspicuous upon sand or mud; its color is, as before, light with fine, dark-red specks. The eyes are black and the chele not red but pink. The liver region is greenish but the dark yolk so long conspicuous has dis- appeared with the perfection of sensory and locomotor organs. Where the head-thorax ends posteriorly there is a dark rim caused by concentration of pigment cells there and the abdomen still has a dark band across the dorsal side of the first somite. The region above the heart is quadrangular and very pale in color. Back of each eye there is a dark longitudinal band. The groove between the head and thorax dorsally is light and the pigment anterior to it is more dense. Under the microscope the arborescent red pigment cells often have a blue background and in some regions there are arborescent yellow cells amongst the red ones. The considerable increase in size in passing from the second to the third stage may be seen by comparing the measurement given above, page 28, with the following. In third stage a larva measured 14 mm. from rostrum tip to edge of telson and 15 mm. to end of plumose sete. The antenna was 12 mm. and the chela 10 mm. long. The width of the thorax was 3.5 mm. and its depth 4mm. The telson was 2 mm. wide and the fan formed by it and the sixth pair of pleopods 5 mm. without the plumose sete, and 6 mm. with them. The length of the head-thorax was 7 mm. and of the abdomen 7 mm. In the third stage the larve were active and voracious, walking and swim- ming with ease and speed so that they were hard to catch. When kept to- gether they soon lost chele in fight with one another and greedily devoured their dead fellows. When a piece of frog’s muscle was put into a dish with these larve they seized as soon as they came into contact with it and holding the main mass with their chele and other claws dragged it backward while tearing off fragments with their mouth appendages. Thus twenty-three to twenty-five days after hatching larve in the third stage which had had no flesh food, unless THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 37 it were the chele of other larvee, possessed well developed responses in the pres- ence of flesh food. When meat was put on green ooze on the bottom of small aquaria containing such larve they showed no sign of being aware of it till in contact with it when they seized it ravenously with their mouth parts and then holding the mass with their chele jerked back like a dog tearing meat from a bone, so strongly as to pull off mouthfuls. When frightened away from the meat, which was then placed one-third of an inch to one side of its former posi- tion in the ooze, the larva returned to the place where the meat had been and seemed to masticate the ooze there and very slowly found the piece of meat again as if through touch, taste, or smell and not at all by sight. The pugnacity of the third larve resulted in loss of limbs and specimens with legs pulled off at the ‘‘breaking joints’? and also rapidly regenerating there were seen. Chantran stated that larve of Astacus would regenerate lost limbs in seventy days while the adult males required one and a half to two vears and the females three or four years. Exeepting the newly expanded last abdominal appendages all of the append- ages agreed with those of the second larva in most all details, but they were larger. The first antenna, however, in place of the eight sense clubs of the second stage, had eleven. These were placed as follows on the exopodite: four in a group on the distal segment; three at the distal end and one upon the middle of the under side of the penultimate segment; two at the distal end of the antopenultimate segment and one at the distal end of the next segment. Just as in the second stage the ear cavity was protected by a row of plumose sete arching over it from its external border. The long filament of the antenna was often broken near the tip but contained from 60 to 65 segments and some few of the terminal ones were constricted about the middle as if they might divide at the next moult. The acicular sete of the filament were about one-half as long as the seg- ment, from the distal ends of which they arose in whorls of five or six. To- ward the tip of the filament these sete were much longer than in the second larva. The mandibles, maxillew, maxillipeds, chelew, and walking legs with their gills and sete were the same as in the second stage except for increase of size. Upon the abdomen the pleopods also were as in the second stage, except in the ease of the expanded sixth pair above described. As yet no appendages were seen upon the first abdominal somite but as in this Astacus no ap- pendages were found there in the adult female it may be that only female larve were examined. The four pleopods of the somite anterior to the sixth still had the exopodite longer than the endopodite, as was also the case in the new sixth pleopods (fig. 42). In the adult this relative size of exopodite and endopodite is reversed in the abdominal organs that serve as secondary reproductive organs. 38 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS Thus in the male the first and the second somites bear the remarkable male sperm conductors in which the endopodites greatly exceed the exopodites or else form the only part developed while in the four following ple- opods the larval relation is not much changed in the adult, as the exopodite is as large as the endopodite or, in the sixth pair, larger, and in all these four the function is not sexual but locomotor, or perhaps respiratory. In the female the same state is found in the sixth pair, the first pair are lacking, while the four other pairs have the endopodite much longer than the exopodite and both are used as reproductive organs, as supports for the eges and larve. Some of the larve of the third stage were kept from May 18 to October 2, and some of the moultings and increase in size noted, but no details of the gradual completion of adult structure were studied. The fourth larval stage was reached by a moult after the larva had lived in the third stage twelve to fourteen days and was a little over one month old. The length then increased from 15 mm. to 17 mm. The color was no longer bright but dull and inconspicuous, pale grey densely spotted with almost black pigment and scarcely any flesh color though the tips of the chele were pink. There was a marked transverse band of bluish across the posterior edge of the carapace and the abdomen was mueh lighter than the thorax. The fourth stage had advaneed beyond the third in one important par- ticular since now for the first time the appendages of the first abdominal somite were seen in some specimens, which were probably males. These pleopods, how- ever, were as vet but very simple rounded knobs which projected from the sternal ridge of the first somite downward and decidedly inward, toward one another, and were about one-tenth of a millimeter long. At the end of June when the larve had been in the fourth stage about a month they passed into a fifth stage which was nineteen millimeters long and in color red-brown or in some eases decidedly bluish. Kept in running water with water plants and tubifex for food and at a temperature as high as 21.5° C., they climbed about actively upon the plants or else remained buried in the ooze. From some of the larve left ina closed aquarium with algal ooze from June 16 to October 2, there remained one survivor five months and a week old that measured 30 mm. in length, 6 mm. in width of thorax, 12 mm. in width of telson- fan, and 25 mm. along the antenna. This crayfish having areas for the ends of the oviducts upon the antepenultimate legs was a female, and it had no append- ages upon the first abdominal segment. The color was bright, finely speckled over with brown; the legs lighter; the antenne dark; the chele purplish with red and blue spots and there was still a blue transverse band across the poste- rior edge of the thorax. The eyes were brown. ‘he shell and flesh were still translucent so that the intestine showed through the dorsal side of the abdomen. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 39 This larva buried itself in the ooze but was so active as to be caught with difficulty. By October 20, it had shed and grown to a length of 34 mm., with antenna 32 mm. long, so that with the food that had been given it it had gained 4 mm. in length in eighteen days. This larva lived till February, 1905, and died when nine months old. Another lot of larve kept in running water from July 16 to October 2, also yielded but one survivor which had increased from less than 26 mm. to 55 mm. and had a telson-fan 25 mm. wide, thorax 13 mm. wide, and antenna 36 mm. long. This fine young crayfish five months and a week old looked like the adult and was a male with well formed male stylets upon its first abdominal somite. The body was opaque and dark; greenish-blue and brown finely speckled; the legs lighter; the chele with much blue and with the conspicuous fiesh-colored area that the adult has at the angle of the claw. The posterior edge of the carapace was dark blue and the antennw were bluish at the base fading to brownish at the tip. The under side of the body was pale flesh-color, or color- less with some clear blue. A third lot of larve, 18 mm. long on May 27, and in the third stage, were kept in more favorable conditions, that is, running water in a large tank, with mud, plants, and sunlight, and on October 2 there were six survivors. The measurements of these larve, five months and a week old are given in the table below: ee ei Ae Ber CG) -D- Rok eine i ann ONOO Go MOSM OO ramo2 Width . es ss ass NG) Ny aM Antennalength . . . S51 52 652 50 55 45 Se xtreereenermach | fate kt g Q fe) co} Q $ The males A and F had well formed male organs upon the first abdominal somite while the four females had no appendages there, but the largest females B and C had two blue spots upon that somite that suggested some connection with a possible appendage. The colors of these large young were those of adults, and though all reared in the same tank they showed individual differ- ences in: general color, some being more dark olive, others brown; in size and color of the spot at the angle of the claw, which in one was reduced to a mere light nodule and in others was a very large area; in color of under side of claw which was dark or else bright flaming-red. The eight young reared to an age of five months and one week thus measured 30, 55, 56, 60, 63, 58, 58, 52 or an average of 54 mm. Tabulating the facts above recorded as to the size and length of duration of the larval stages as far as they were followed we find the following: 4() THE YOUNG OF THE CRAYIFISHES ASTACUS AND CAMBARUS Stage. Duration. Lene of Habit. 2 ody. 1 | 4-138 days 9mm. | Fast to parent. Dy Bagh & 11 mm. | Free and gradually becom- | | ing independent. | Se pl2=14 14-15 mm. | Independent. | d aio) 17 mm. | Independent. 5 2 19-20 mm. | Independent. The number of moults necessary to increase the larva from the fifth stage when about 20 mm. long to the autumnal larve ranging from 30 to 63 and averaging 54 is not known. The amount of increase at each moult above tabulated was about 3 mm. In one case above cited a larva which had grown only to 30 mm. in the autumn, when well fed quickly gained 4 mm. probably in one moult. We might expect then an increment of 3 or 4 mm. at each moult, and growth from the 20 mm. length of the fifth stage to the average autumnal length of 54 mm. may have taken from 8 to 11 moults thus making an estimated total of 12 to 15 moults the first growing season. The size reached was, how- ever, evidently determined in part by food. On such basis it may have taken even more moults to produce the large autumn young 60 and 63 mm. long. And at all events the above larva 30 mm. long having been kept in a closed aquarium should be disregarded in reckoning the average. Rejecting this the average for the seven others reared in running water would be 57. To attain this average size it may well have required at least nine moults after the fifth stage. It seems then not improbable that this Astacus, under these conditions moulted at least a dozen times while growing to a length of over two inches in the first five months of its life. Four stageswere observed in the first two months when the length had not extended to four-fifths of an inch, and probably twice as many stages were necessary in the following three months to bring the length up to over two inches. The only previous records of the rate of growth of young Astacus seem to be those of Soubeiran, Chantran, and Steffenberg. Soubeiran (765) from measurements of a crayfish in a French crayfish farm concluded that they did not moult more than once in the first year, and were 50 mm. long when one year old. Chantran (’70) thought the young crayfish moulted five times in 85 to 100 days of July, August, and September and no more till the end of the next April. The first moult was at ten days after hatching and the other four at in- tervals of twenty to twenty-five days each. These results were modified by his further studies, also in the laboratory, and later (’71) he stated the number of moults in the first summer was eight and that the temperature influenced the THE YOUNG OF THE CRAYTFISHES ASTACUS AND CAMBARUS 41 number of moults so that there were six in the second summer, if hot. Finally Steffenberg (’72) gave the lengths of the Astacus larve in Sweden as 8.5 mm. at hatching; 11 mm. in the second stage; 13 mm. in the third stage; 15 mm. in the fourth stage. Astacus leniusculus thus agrees closely with the Astaecus of Sweden in the length of the larva in the first, second, third, and fourth stages. As far as the facts go it seems evident that Astacus leniusculus probably has mare larval stages than have been described for the first year young of the French Astacus, but the differences are probably due more to difference in food and in temperature than to any innate differences in the species. With this close agreement in larval life between the American and Euro- pean Astacus and the demonstrated success of rearing in the laboratory the large young from the eggs brought overland, the culture of the American As- tacus should be as successful when undertaken as has been crayfish culture in Europe. As elsewhere remarked (Andrews, :06°; :06*) the introduction of the western Astacus leniusculus into Eastern waters might not only prove of economie value but also help to throw light upon the interesting problem of the nature of the causes that have brought about the present remarkable geograph- ical distribution of crayfish. And the geographical distribution of crayfish is intimately connected with the origin of species in this group. CAMBARUS AFFINIS. As elsewhere described (Andrews, :04+) this common crayfish of Maryland and adjacent States lays its eggs in the spring, and the development of the young can be followed in the laboratory. Preparatory to laying, the females earefully cleanse the parts of their bodies to which the eggs are to be attached and the eggs flow out of the oviducts into a mucous mass which covers the ple- opods upon which, after some special ‘‘turning’’? movements of the female, the eggs are found attached each by its own stalk (Andrews, :06). The hatching young thus find themselves upon the abdomen of the parent and here, as in Astacus, there are special contrivances which prolong the connec- tion of parent and offspring for some time after hatching so that the attainment of a free and independent existence is a slow and gradual process. The special arrangements used in the attachment of the egg and of the larvee in the first and second stages form a series of interesting adjustments between the adult and the next generation. These successive means of association of mother and offspring will be described in what follows. We will first consider the eve and the maternal appendages. The eggs, while numerous, three to six hundred according to the size of the female, are small, scarcely 1.5 mm. as compared with 2.5 mm. in Astacus len- iusculus. Some few become fastened to sete upon the abdominal sterna but 4 42 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS most are fastened to plumose hairs upon the pleopods of the second to fifth somites inclusive, and some account of these pleopods may be given here in connection with questions as to how the eggs and larve are attached to the mother. The posterior face of the fourth left pleopod of a female about to lay is represented in figure 44. It will be noted that the endopodite is longer than the exopodite, and both bear a fringe of long plumose set on their right and left edges. The sete are shorter toward the base of the exopodite and of the endopodite while on the protopodite there are but a few plumose sete in two tufts upon the inner edge. The long sete make of the pleopod a wide fan or flat brush since the sete lie close together and like the wing feathers of a bird form a rather flat firm expanse. The protopodite has a basal part containing several calcified plates in a soft membrane, and a long segment that is well caleified except fcr a triangular soft area toward its base on the posterior face. The endopodite is made of two segments and the exopodite of one. The endopo- dite and the exopodite are also somewhat annulated in appearance owing to the grouping of the cement glands. The groups are opaque white and from each side tend to run together across the posterior face. Distally they do not meet but proximally they meet and make cross bands. Still farther toward the base the glands cover the entire surface more and more completely. The non-gland- ular areas are clear and not opaque, and in the figure are represented dark. The anterior face differs from the posterior (fig. 44) in a greater de- velopment of glands which formed transverse bands more nearly all the way to the tip. As so many of the glands are to the right and left near the setw, they are well placed to smear their secretion over the sete. Toward the tips of the pleopods the exoskeleton is so translucent that with Zeiss 2 D, the striation of the muscle fibers, the branched connective tissue cells, granular blood corpuscles, and the polygonal gland cells may be seen. The gland cells are about the diameter of a muscle fiber and larger than a blood corpusele. In most cases the plumose sete spring from over the glands, and the base of a seta is as thick as two gland cells. The set have a large central cavity and a thick wall which is highly refractive and clear and is of unequal thick- ness so that it projects into the cavity in lumps or waves, and gives the distal part of the axis of the seta a somewhat segmented aspect. At the base each seta is articulated to the exoskeleton, and its central cavity is constricted by a clear refractive thickening of the wall that leaves very little communication be- tween the cavity of the set and the cavity of the body. The side branches of the seta spring out not only along its sides but also, seatteringly, along its posterior face so that the plumose seta is more like a bottle brush than lke a flat feather. While the side branches generally make THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 43 a wide angle with the main axis, toward the tip they become more nearly par- allel with it and thus a fine terminal brush is formed by the main axis coming gradually to an acute point in the midst of the surrounding side branches. The eggs are fastened to these plumose hairs by a secretion that probably comes out of the glands of the pleopods (Andrews, :06°). They then look as in figure 45, which represents the anterior face of a pleopod cut off in the afternoon of April 18, 1905, from a female that had laid the night before. Most of the pleopod is concealed by the eggs which are opaque yellow balls and very elastic. The plumose setx are all bound together by a common cement or mass of glaire so that the individual sete are not seen. From this mass a clear, flat, glassy band of material goes out to each egg. Upon separating the eggs these bands are seen as clear, flat stalks, continuous at one end with the mass that binds the sete together and at the other end with the envelope about the egg. While the shorter bands are flat and wide the longer bands are more string-like and some few are twisted. Though these stalks may cross one another and be more or less intertwined they are not fastened to one another. As the sete spring chiefly from the side of the pleopod and the eggs are tied to the sete, the eggs may be combed out, as it were, into groups on each side of the endopodite and of the exopodite. Where the stalk joins the egg it is enlarged as a bell, or tent, full of liquid and its edges are continuous with the outer layer of the egg capsule. Each egg had then its own separate stalk, though a very few exceptions showed two stalks connected with an egg at different points and with a com- mon extension running over the surface of the egg from one stalk to the other. Thus very firmly attached to the mother, the eggs are waved back and forth by the mother who regulates the movements of the pleopods in accordance with the oxygen supply in the water, and thus they slowly develop till they hatch. The old egg cases and stalks still remain fast to the pleopods, and are of use to the ‘larva as a means of prolonging its life of dependence upon the mother. 2 We will next describe the way in which the first larva is connected with the mother. In hatching, the larva comes slowly out of the egg capsule through a rent along its back, in such a way as to draw out the legs and abdomen last of all as represented in figure 8 in a previous paper (Andrews, :04). In fact the tip of the abdomen remains inside the egg ease long after the soft, helpless larva is extruded and left dangling down into the water. And all these newly hatched larve would fall to the bottom were it not for a firm attachment of the tip of the abdomen inside the egg case. As it is some time before the respiratory move- ments become perfect, as the limbs only gradually acquire ability to move, and as the body is globose and the creature cannot stand on its legs, the larva would 44 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS probably perish but for this temporary fastening of the abdomen that tides it over the weak period till it can reach up and take hold of the egg stalk as in- dicated in figure 9 of the above paper. This attachment of the abdomen is the same phenomenon found in Astacus and is brought about by a telson thread; but as the eggs and larve of Cambarus are so much smaller and as the tip of the abdomen remains inside the egg cap- sule the facts are not so readily made out and in a previous notice (Andrews, 04) the telson thread was spoken of as proceeding from the anal region, though further study shows it to be fastened to glandular spines of the telson edge as in Astacus. In Cambarus affinis the tip of the abdomen is fastened by a short thread to a erumpled membrane which lies inside of the spheroidal egg case and is firmly fixed to the egg case on the side near the stalk. The short thread and the membrane together are comparable to the long membranous thread that im Astaeus allows the larva to hang far down from the egg case. For conven- ience we will describe the two parts of the telson thread separately. The short part fastened to the abdomen remains on larve 48 hours old though then broken off from the more membranous part inside the egg case. The short part may be whipped up and down in the water like a lash when the larva flaps its abdo- men (fig. 50). At hatching, the telson (fig. 46) is a simple rounded, translucent lobe, with minute spines on each side of the median plane, which formerly fastened it to the telson thread. In this ventral view of a recently hatched larva the pleopods of the fourth and fifth somites are seen free while the sixth somite has its pleo- pods as lobes inside the telson on each side of the anus. The terminal part of the telson is traversed by radiating lines which point to spines along the edge of the telson. These lines are in reality rows of cells that are to make the plumose sete of later larval stages and the rest of the translucent flat telson is filled by a parenchymatous mass traversed by blood spaces in which float blood corpuscles. The telson thread arising from the edge of the telson is a flat band that is readily twisted and shows a striation due to fine wrinkles of the stiff chitin-like material composing this very strong but thin and translucent membrane. Twenty hours after hatching, the telson had changed form, become more quadrangular and its terminal part was somewhat three lobed (fig. 47). And as the spines to which the telson thread was attached were all on the middle lobe it seemed as if the pull of the telson thread might have aided in making the ~ middle region protrude as a lobe. At this time the cuticle of the larva was separating from the body in preparation for the moult from the first to the sec- ond stage. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 45 While the form of the telson is thus so different from that of Astacus (fig. 90), the mode of attachment of the telson thread is fundamentally the same. Figure 48 shows the tip of the telson of a larva torn out of its shell just before hatching; the thread is fastened to five or six spines on each side of the median plane in the same way as in Astacus (fig. 21). Thus nearly the same number of spines are specialized in both crayfish for attachment of the thread though in Cambarus affinis the entire number of spines is much less and they are found only on the posterior part of the edge of the telson. All the spines are glass clear, ice-like in refraction; the lateral ones are bent toward the tip of the telson and the five or six specialized glandular spines converge toward a center as shown in figure 48. The lateral spines toward the posterior end tend to show blunt brushes and_ secreted lobes at their tips, and thus form somewhat of a transition to the effective specialized spines. These specialized central spines are much longer and thicker and bent, often at right angles. Some are fused together by their blunt ends and all seem to have flowed out at the tips as a mass which is now fibrous and which binds all of them to each other and to the telson thread. They seem compar- able to paste tubes which should squeeze out a myelin-like substance that could coagulate as strong fibrils. The appearances suggested that the rows of gland cells that later make the plumose sete of the later larva, had previously, in late embryonic life, seereted a substance which oozed out of the hollow spines and set into a firm cement; but these cells no doubt were also active in making the cuticular walls of the spines themselves, and no sharp line seemed drawn between the substance of the cuticular spines and the material that issued out of their tips. Both are pre- sumably the same exoskeleton and made from ectoderm cells that later make other exoskeletal secretions in the form of plumose sete. In Astacus (fig. 21) the distinction between spine and secretion was more evident, but in this Cam- barus (fig. 48) the spines are so minute that details are not as readily seen. While the mode of attachment of the telson thread to the telson is thus the same in Astacus and in Cambarus, the thread itself differs in appearance in the two crayfish, in the former being pulled out into a long thread, in the latter being for the most part a wrinkled mass of membrane within the egg ease. While in Astacus the thread is apparently a cast off embryonic skin, this is by no means obvious in Cambarus and an interpretation of its meaning was had only from the following facts. When an egg ready to hatch was seratched with a needle the outer egg case came off and the larva popped out alive but still enclosed in a thin spheroidal membrane. This membrane was firmly fastened to the outer egg case by one small area towards which the legs con- verged and which lay opposite the claws. When the egg case was pulled it re- mained so fast to the membrane that both were drawn out of shape rather than 46 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS separate. The part of the egg case attached to the membrane was often near to the stalk of the egg case, which led one to infer that the attachment of egg case and membrane and the making of the egg stalk might have some common cause very early in the life of the egg, but no observations were made to de- cide this. Left to itself a few minutes, such an embryo removed from the shell burst the membrane without the aid of the egg case. Its telson was fastened by its special glandular spines to the inside of the membrane, and soon it reached about with its claws and seized hold of the membrane. This bag-like membrane was thus the same thing as the telson thread of normal hatching. When dissected from an embryo this membrane was seen to be a bag fast to the telson spines, but with no observed special envelopes for the limbs as would be the case were it a cast skin. Before hatching, the membrane was stretched tightly all over the abdomen except where the two special groups of glandular telson spines were, and there the membrane was raised up as two swellings, one right and one left, much higher than the spines themselves. The space be- tween the membrane and the spines was occupied by a mass of seemingly liquid lumps which in some cases with 4 D showed a finely fibrous material amonest the clear spines which, in places, extended out to the membrane like a fibrous coagulum binding the spines to the membrane. On the median line the membrane was close down against the telson so that the secreted masses right and left seemed to have locally pushed the membrane away from its original connection with the telson. This membrane was well developed in embryos of stages J to IK of Reich- enbach but upon dissecting, these embryos dropped out of the membrane, as the telson was then not yet fast to the membrane, though the membrane was firmly fastened to the egg case near the stalk. In the early stage F there was also a membrane over the body and this was loose over the slightly projecting, small abdomen. It seems probable that the telson thread of Astaeus, which is a cuticle formed over the embryo when its limbs are well advanced and thus has tubular outgrowths to cover the limbs, is represented in Cambarus by a telson thread having the form of a sac-like membrane formed so early that the small limbs receive no special envelopes. In both eases there is a special outgrowth to cover the abdomen, but while in Astacus this is a long bag, in Cambarus it is searecely recognizable as a separate region. In both, by some unknown process that is imagined to be associated with fertilization phenomena, the membrane is made fast to the outer eg@ case, and in both the membrane becomes fastened to the embryo by the activity of certain telson glands. A diagram of Astacus would represent it as escaping from a cuticle when hatching, a euticle fastened to the eg@ case at one point, and near that point fastened, inside, to the telson of the larva. A diagram of Cambarus THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 47 would represent it as casting off a membrane at hatching which is fastened as in Astacus but lacks the tubular coverings for the limbs. Here again the small - size of Cambarus may have led to misinterpretation of what is plain in As- tacus. Having described this temporary, mechanical association of parent and young we will next consider the more active association that lasts during the rest of the first larval period. The larve, supported for a time by the telson thread, soon established a second connection with the mother by seizing hold: of the ege case, the egg stalk, or in many cases the matted pleopod sete, with their chelew, and thus, for a time, were fastened both by the telson thread and by the chelw. While the eggs hung loosely from the pleopod, figure 45, the young erayfish had the habit of reaching in their chele as far as possible amongst the egg stalks and pulling themselves close to the pleopods so that they beeame densely crowded together in a solid mass (fig. 52). In life this mass of young cover- ing the pleopod is a curious sight suggesting mammalian young crowding for maternal milk. Each held itself close to the pleopod, and when disturbed drew itself all the closer as if eager to remain. The part exposed to view is chiefly the rounded head-thorax which is flesh-colored but is conspicuously marked by the enclosed saddle-shaped dark yolk. Here and there the legs and abdomen are seen in profile (fig. 52), but generally the abdomen and legs are under the body, the abdomen being bent somewhat as ina brachyuran (fig. 50) and a full dorsal view shows chiefly the simple head-thorax (fig. 49). The larvee thus seem egg-like, inert and inactive. The long-stalked egg cases stand out above the backs of the larve as do also a few belated eggs which may hatch some forty-eight hours after the rest (fig. 52). By this time many of the larve have broken the telson thread and when disturbed make slight movements with their legs and flap the abdomen to which is fastened, like a small handkerchief, the narrow telson end of the thread, while the membranous sae remains within the egg case. , So closely do the larve crowd together that only few of the pleopod setie can be seen. Figure 52 represents a case in which some of the larvee had been removed. While the color of the first larva is light flesh-color to the naked eye, under the microscope the creature is clear, and colorless, except for the scattered, arborescent, vermilion pigment cells over the carapace, abdomen, basal parts of antenne and some few segments of the legs, and except for the reddish pig- ment in the eyes and for the large red-brown yolk mass. The shape of the larva when alive is retained very well in specimens killed in Woreester’s liquid, though in many other liquids the head-thorax swells and is abnormally glubose, while the branchiostegites roll back and expose the gills. 48 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS In neither the dorsal nor the side view (figs. 49, 50) can the rostrum be well seen, but in a front view (fig. 51) it is seen to be pointed, though so bent downward be- tween the eyes as to be of no use as a protection. The body is thus rounded and embryonic in proportion and lacks all the angularities of adults or of active larve. With the clinging helpless state of the larve-is associated a lack of ordinary use of the antenne. The second antenne, which in larval and adult life of crus- tacea are carried out in front of the animal, are here folded down under the thorax (fig. 50), between the right and left series of legs. This peculiar posi- tion of the antenne is not found in Astaecus and here is brought about at the time of hatching, for in the embryo these antenne grow backward along the edge of the carapace external to the bases of the limbs. Soon after hatching the legs get astride the antenne and this apparently useless position of the antenne is maintained during the rest of the life of the larva in its first stage. This peculiar position of the antenne as well as the down bent rostrum were observed in another species of Cambarus, C. rusticus by Faxon (785) ; and later Steele (:02) described the rostrum as bent down in C. gracilis and in other species and the legs, antenne and abdomen of C. virilis as lying under the thorax; thus it seems possible that the first stage of Cambarus in general differs from Astacus in these characteristics of the antenna. The appendages of C. affinis in this first stage resemble those of Astacus leniusculus in being without the sete of later stages, but they differ not only in being very much smaller but in being, in some cases, more simple. Thus the first antenna (fig. 53) has only four segments in the endopodite and in the ex- opodite in place of five, and the tip of the endopodite bears no sensory club. The antenna (fig. 54) is remarkably short, as is seen on comparing that figure with figure 7, and figure 50 with figure 3. The filament is bent back and has but twenty-five segments, or about half as many as Astacus. In the adult there may be 150 segments. Carried as it is under the thorax it reaches only to the beginning of the abdomen while in Astacus the antenna if in such a posi- tion would reach about to the end of the abdomen when stretched out. The mandible (fig. 55) has the same simple form as in Astacus (fig. 8). The first maxilla (fig. 56) is like that of Astacus (fig. 9), but more simple in lacking the few filose sete and in having fewer spines. The second maxilla (fig. 57) as in Astacus (fig. 10), has a row of plumose sete along the entire edge of the scaphognathite and is elsewhere as simple. The maxillipeds (figs. 58, 59, 60) represent in miniature the structure seen in Astaeus (figs. 11, 12, 13) with shgeht increase in smoothness due to the pres- ence of fewer sete and spines. There are important simplifications, however, in the gills which in some cases have but half as many side filaments as in Astacus. The chele (figs. 61, 62) have the same recurved tips as in Astacus. Soon THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 49 after hatching the larva opens its claws widely (fig. 50), and after some failures fastens them to the egg stalk, or to the setw# of the mother’s pleopods. Once firmly locked in the hardened secretion that makes the egg stalks and binds the sete together, these larval chelwe remain fast during the first stage. And even after the larva has escaped from its cuticle and passed into the second stage the old cuticles are left firmly hung to the mother by the cast-off chele. As in Astacus the chela has at first a cutting edge set with a row of few and simple spines, but when the first larva is ready to moult, the loosening of the cuticle reveals the fact that each spine will be replaced by one that is ser- rated (fig. 62), owing to the presence of flat plates along the posterior face of the new spines. In addition the second stage will have on its claw some spines not represented in the first stage and also near the tips of the claw some long, simple sete in place of the blunt spines there in the first stage. This figure shows that the recurved tips will be abandoned in the second stage since there are already formed tips that are but slightly curved hooks, to take the place of former reeurved tips, one of which was broken off in this specimen. The chela besides being so small and weak, is inferior to that of Astacus in having its gills less developed, the anterior arthrobranch being very short and simple and with but few side filaments. In the four walking legs (figs. 63, 64, 65, 66), we find the same proportions as in Astacus but the pleurobranchie are absent and the arthrobranchiw are more simple, especially the anterior ones. The branchial formula for the first stage is thus the same as in the adult and is as follows: Arthrobranchiz. | | Podo- = Pleuro- | Total branchie. | branchie. | ean Anterior. | Posterior. | z —— | = | | Somite of 2d maxilliped . . . . «| 1 1 0 0 2 Somite of 38d maxilliped 1 1 1 0 3 Somite of chela ; 1 1 1 0 | 3 Somite of Ist walkinglez. . . . . 1 | 1 | 1 | 0 | 3 Somite of 2d walking leg. . . - .-| 1 1 | 1 | 0 3 Somite of 8d walking leg. - . . -| 1 1 | 1 0 | 3 Somite of 4th walking leg . . . -| 0 | 0 0 0 0 ——— | —————— 6 | 6 | 5) 0 | 17 That the pleurobranch of the last thoracic somite of Astacus was absent from Cambarus rusticus larve 4 mm. long and evidently just out of the egg was observed by Faxon (’85) and it is probable that no members of the genus Cambarus have remnants left in the early larve of that pleurobranch still found in Astacus. 50 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS As in Astacus the abdomen in the first stage of C. affinis, just as in some other species of Cambarus studied by Faxon (’85), has only four pairs of evi- dent pleopods. The sixth pair are but partly formed and enclosed within the tel- son while the first pair exist here as minute buds readily overlooked and not found in Astacus nor as yet described in other species of Cambarus. The four evident pleopods (fig. 67) have equal exopodites and endopodites in contrast to the proportion found in Astacus (fig. 19). The beginnings of the first pair of pleopods are but minute rounded elevations of the sternal ridge of the first somite (as indicated in fig. 8; Andrews, :06), and contain a small mass of nu- cleated epidermal cells. The appendages of the sixth somite are large internal buds that cause the side part of the telson to be very thick as they fill all its interior on each side of the anus. As this region is very translucent each pleopod ean be seen to have a long outer and a short inner lobe (figs. 46, 47). The first larva is in several ways more imperfect than the same stage in Astaeus, and as it lives inactive and fastened to the maternal pleopods it seems but an extension of the embryonic period in preparation for the second stage, and not at all a self supporting organism. The second stage is made way for by the gradual loosening of the cuticle of the first stage, as above noted in case of the claws of the chela, and by the addition within the old cuticle of the new exoskeletal structure of the second stage. This was evident in the telson of a larva twenty hours after hatching, where the old cuticle was raised up the length of the old telson spines and new spines had grown out across this space to enter the hollows of the old spines. Later, just before the moult, the space between the old and new cuticle was twice the length of the old spines, which were still fastened to the remnant of the telson thread, and the new spines still projected across the space into the old hollow spines. The moulting into the second stage took place after the first stage had lived this dependent and preparatory life for about forty-eight hours and the larva resulting had the form shown in figures 68 and 69. These second larve unlike the second stage of Astacus did not gradually become separated from the mother and live isolated, but remained again adhering to the pleopods till a second moult brought them into the third stage which was the first free larval form. Thus the pleopods were still covered with the crowded larve. Figure 70 represents a pleopod with only part of its load of second stage larve as many had been removed. . In the following description it will be shown that in the second larva also there is a peculiar temporary mechanical connection of young and parent. Each pleopod in life was covered as with a mass of animated jewels of pink and garnet colors. The flesh-colored larve were still strongly marked by the yolk that remained in each as a garnet red mass of saddle-bag shape. No longer firmly fixed by the chelw, the larva soon reached about for firm objects to THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 51 which to hold, or kicking their legs about, swinging their pleopods and _ flap- ping their abdomens formed an animated restless mass quite unlike the inert one seen in the previous period. Thus active, though holding fast, the larve were no longer densely crowded in close to the pleopods as in the first stage (fig. 52), but loosely aggregated (fig. 70). The old larval cases shrivelled and not readily recognizable as more than erumpled membranes, still remained fast by their chele and the old egg cases were yet fixed by their stalks to the maternal pleopods. Side by side on old ege stalks were the pairs of living claws and the empty cuticles of old cast off claws. Thus the second larve lived amidst a mass of former envelopes,—the ghost-like phantoms of their former selves and the rigid cradles of those selves. - While many larve holding fast to the pleopod hairs and to egg stalks tend to pull themselves in close to the pleopod so that they become buried amongst the ege stalks and empty cases, a few larve at the periphery of the mass may dangle free in the water, as in the lower right corner of figure 70. These are larve which have recently cast off the first cuticle and are still in a soft, help- less condition. Before these larve succeed in reaching up and seizing hold of some part of the general mass they are in danger of falling to the bottom where they might be lost, though when they are removed it is found that they can stand and walk, with difficulty. However they seem strongly controlled by an instinet to climb, crawling over one another in heaps when removed from the mother and swimming only when thrown into Perenyi’s liquid and possibly many would eventually climb up onto the mother. At all events they are prevented from fall- ing away from the mother by a fine thread which, as indicated in figure 70, passes from the posterior end of the larva to the crumpled, cast-off cuticle which still is firmly fixed to the mother. This thread may be called the anal thread since it actually comes from the anus and not from the rim of the telson as did the supporting thread of the first larva. But before discussing this anal thread some general features of the second larva may be considered. The abdomen is still carried bent down as seen in figures 69 and 68, but the cephalothorax is more elongated and narrow than in the first stage and ends in a sharp pointed toothed rostrum that projects out be- tween the eyes and is only slightly bent down. The legs are longer and may now be used for walking and the antennx stand out in front of the animal as if use- ful feelers, in strong contrast to their apparently useless position in the first larva (fig. 50). This increase in length and change of position of the antenne gives the mass of young in the second stage (fig. 70) a much more living aspect than was possible in the first stage (fig. 52). The abdomen when straightened out is seen to end still as a simple rounded telson (fig. 71) with the large sixth pleopod still within it on each side the anus and with a terminal part that has its middle lobe set off but slightly from the side lobes. The edge of the telson 52 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS bears only simple spines as in the first stage (fig. 46), and these spines are not divided into groups (fig. 72), but all seem equally useless since none act as glands and there is no telson thread formed in the second stage. The middle lobe bears spines all along its edge, some sixteen on each side replacing the fewer spines of the first stage (fig. 48). Passing in from each spine there is still a radiating line of cells which is active in making the plumose sete which we miss in this second stage but which will be expanded in the third stage. It is to the larva of the above structure that the anal thread is fastened and this may now be considered in connection with this telson that has pro- gressed so little beyond its first form. As seen in figure 71, the anal thread ras in life a clear narrow ribbon running from the anus a distance greater than the length of the telson to be fast toa crumpled mass which proved to be the cast-off cuticle. This thread was fast at one end inside the intestine of the larva and at the other was fast to the cast euticle by being continuous with it at the edge of the cast off anal opening. In fact the anal thread is nothing more nor less than the old cuticular lining of the intestine of the first larva still con- tinuous with the cast-off cuticle of the abdomen and not yet entirely loosened from the intestine of the second larva. An examination of the recently shed larva showed that the cuticle of the larva had become loose all over the body and some distance into the intestine, but that farther in it still adhered to the epithelial lining of the intestine. When the larva has moulted, the old cuticle of the abdomen is found to be telescoped and its old telson joined by a short cord to the new telson as shown in figure 73; a. If the old euticle is then pulled with forceps it does not break loose but the thread comes out of the anus of the larva and is thus made longer, as shown in figure 73 8, by a shortening and crumpling of the intestine. There is thus a posterior part of the intestine (y) in which the old cuticle is loose and an anterior part (7) where it is still fast, the region (a) being the first place met with where the cuticle lining of the intestine does not pull away from the in- testinal wall. Hence the region (y) is thrown into folds and the region (2) stretched and pulled bodily toward the anus when tension of the anal string pulls the region (@) toward the anus. The anal thread is the loose lining of the intestine as far as it is pulled out of the anus and while in origin a tubular cuticle it is stretched out as a flat ribbon which seems made of clear wire-like fibres, but in reality is only thrown into longitudinal folds due in part to the longitudinal ridges which sections show are present in the intestine. In figure 74 is seen more enlarged the region of the intestine where the loosened cuticle running up from the anus arrives at the region (a) where it is still fast to the epithelial walls, as made out in preparations cleared in Bela Haller’s liquid. Though actual ecdysis was not seen, it is evident that what happens in the moulting of the first larva to the second stage, is that the lining of the intestine THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS d3 for a certain distance up from the anus, delays in casting off its cuticle. The animal bursting out of its head-thorax cuticle and freeing its limbs from their cases would next pull its abdomen out of its old euticle, but the old cuticular lining of the intestine remains firm where continuous with the outside cuticle at the edges of the anus, and acts like a string tied to the old telson and to the inside of the abdomen far up in the intestine. If the larva then drops out of its cuticle and is too weak to seize hold of the old eutiele, or neighboring firm objects, it will dangle suspended by the anal thread and the tension on this thread will pull it out of the anus as far as possible. By puckering the posterior part of the intestine and by dragging the anterior part of the intestine backward there is thus drawn out of the anus an anal string which, however, is not as long as the abdomen, by any means. The old telson being a stiff plate and the old walls of the abdomen being elsewhere like the bellows of a camera, the pull of the weight of the larva along the telson thread drags the old telson against the cast-off abdominal rings and telescopes the abdominal cuticle as if the front board of a camera were pulled back against the bellows by a string inside. The tendency of the telson string would be to turn the abdominal euticle of the cast- off case inside out as a hand holding fast to a sleeve would turn the sleeve inside out when the arm was withdrawn, but the presence of a firm, board-like telson at the bottom of the sleeve allows only a telescoping of the cuticle. By this means the old abdominal walls are so shut up that the free anal string from the old anus to new anus is no longer inside the old abdomen, but largely free in the water, as indicated in the diagram on page 54. Soon after hatching, presumably, the cuticular lining of the intestine be- comes loosened along the regions (a) and (#) and then the anal thread is pulled out from the intestine as is usual in moulting. In the meantime the larva becomes strong enough to reach about and take hold of the old larval skin or of other objects fastened to the pleopods and does not need the anal string as a means of support. By this delay in the casting off of part of the cuticular lining of the in- testine an advantage to the larva seems to result. Both in the first stage and in the second stage the larva has a mechanical means of fixation to its mother during the brief period when it is unable to use its claws for this purpose. Reviewing the life of the crayfish up to the third stage with the aid of a rude diagram (page 54), with reference to the association of mother and offspring, we have seen: the egg fastened to the pleopod set by a material that hardens soon after the egg is laid; the larva hung for awhile by its telson thread; then holding by its claws as well as telson thread till the latter breaks when claws alone hold it locked to the mother; then dangled loose as a helpless second stage supported by an anal thread till able to take hold with its claws and cast off the thread, that is, to finish the moulting; finally moulting again and as an active 54 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS third larva living no longer mechanically bound to the mother, with whom it gradually ceases to associate. The mechanical supports, the telson thread and the anal thread, are left hanging with the egg stalks, egg cases and the other part of the cast off membrane and cuticle; all fast to the pleopod sete which are bound together in a secreted mass. All this material soon disappears, apparently being eaten up by the third larva before it leaves the mother to hunt other food in wider fields. EXPLANATION OF DIAGRAM. Pl. =pleopod of mother; Pl. H=setze of pleopod ; 1=cast cuticle of first larva; St.=egg stalk . Sh. =egg shell; M.=membrane inside shell; Tf.=telson thread, which is broken; Af.=anal thread ; =second stage on the mother’s pleopods held by anal thread till the claws have taken hold. Of the two threads which we here describe as mechanical means for pre- serving the association of the mother and offspring and prolonging the ma- ternal protection beyond the egg stage over larval periods that are especially helpless and not self supporting, the first, the telson thread, is a secretion of glands active before the larva hatches combined with an adherence of the larval envelope to the egg shell that was determined very early in embryonic life. The second thread, the anal thread, is a much more simple and temporary attachment brought about by a delay in casting off the cuticular lining of the intestine and thus utilizing what is cast off as a means of keeping the larva fastened to its old cuticle till the claws can lay hold of some firm object. When the second larva finally becomes fastened by its claws it remains some six days in a condition of little activity. It is dependent upon the mother for physical support though in a state intermediate between the invariably fixed first larva and the wandering third larva. The shape of the claws (fig. 84) shows that they would not become as firmly locked as did the claws of the first larva (figs. 61, 62), though they might stick firmly into soft material. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS aD When larve in the second stage were put into Kleienberg’s picrosulphuric they flapped their abdomens for some minutes but did not loosen their claws from the pleopod of the mother; when put into alcohol, however, they dropped off, and when into Worcester’s liquid they all broke loose and fell to the bottom where they lay kicking their legs for some time as they were not so readily killed as were the third larve. The second larva lives much as did the first larva and in structure is very like it but it differs not only in the above mentioned characteristics of the telson and mode of fixation to the parent but also in the size, proportions, and amount of armament with sete of some of the appendages as will be seen when the nineteen pairs of appendages are taken up in sequence. The first antenna (fig. 76) has grown much longer than it was in the first stage (fig. 55), and the basal segment of the exopodite has divided into two so that there are now five in place of four segments. And as the figure shows there are now a few sete upon endopodite, exopodite, and protopodite and of these there are five blunt sensory clubs on the inner face of the exopodite. Two of these are close together on the distal end of the fourth segment, two are close together on the swollen basal half of the fifth and the remaining one stands alone on the narrow terminal half of same segment. The ear is still a simple deep pit without as yet any sete along its edges but with only seven or eight spines or teeth seen with 2 D along its external edge where the second stage of Astacus has plumose sete. The second antenna (fig. 77) has greatly grown in length and in perfection of form and as above noted (fig. 69) is no longer carried in the remarkable position it occupied in the first stage (fig. 50). Yet while in a position to be of use as a sense organ the antenna is still markedly lacking in sete: the exopodite seale bears only a row of few spines and the long endopodite bears but a few acicular sete toward its tip. With its increase in length there has been also an increase in number of segments in the filament which has now 36 beyond the three large basal segments in place of the 22 of the first larva. The mandible (fig. 78) has added a row of small acicular sete along the medial face of the end segment of the palpus, and developed sharp teeth along the heavy cutting edge of its base. When the palpus is folded down into the hol- lowed face of the mandible two teeth are dorsal and five ventral to the tip of the palpus. The first maxilla (fig. 79) has changed but little, but its spines are longer and sharper and it has developed a few, minute, acicular sete. This stationary state is still more pronounced in the case of the second maxilla (fig. 80), which has only grown sharp sete in place of the blunt termi- nal spines. The changes in the first maxilliped are also the development of sete in place of spines and the addition of a few sete (fig. 81). 56 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS In this second maxilliped (fig. 82) this substitution and addition of set is more pronounced but there is a noticeable retrogression in the gill which is more simple, no longer having any lateral filaments. The third maxilliped (fig. 83) again, as compared with the first stage (fig. 60), has quite long acicular sete in place of blunt spines, and a few sparsely plumose set, noticeably a group at the tip of the exopodite. While the pos- terior gill is longer the anterior is here also somewhat reduced. The chela now used as before for clinging to the mother has its tips some- what recurved (fig. 84), but they are much straighter than in the more firmly fixed first stage (fig. 61). There are added a few acicular sete and the claw is more like the adult in the development of rasp-like spines along its edges in place of the simple spines of the first stage which, as seen in figure 62, were being replaced during the first stage by the toothed spines of the second stage. The four walking legs (figs. 85, 86,87, 88) have changed chiefly in the ad- dition of a few terminal sete and of a few exopodite, thread-like sete. The branchial formula is thus the same in the second as in the first stage. On the abdomen the four functional pairs of pleopods (fig. 89) are now long and slender but very simple and not yet fringed with sete, though the spinules on the edges of the exopodite and endopodite are more numerous than in the first stage. The appendages of the first abdominal somite exist as yet only in the form of minute round knobs (fig. 4; Andrews, :06), but slightly larger than in the first stage. The appendages of the sixth somite are still within the telson (fig. 71). When the larva is nearly ready to moult into a third stage these pleopods make the telson protrude laterally more than in the above figure through the sub- stance of the telson, allowing one to see the form both of the pleopods and their sete. When they are expanded, at moulting into the third stage, these pleopods have the appearance shown in figure 90. The second larva remained fastened to the mother for six days, appar- ently eating nothing, and as the yolk-mass gradually diminished it seems probable that the larva still subsisted upon the original supply of energy taken from the ovary as yolk. Before the moult into the third stage the gas- troliths became quite conspicuous as blue areas showing through the body on each side of the stomach. The actual moult occupied but a few minutes and as usual the head-thorax came out of the old shell first, then the legs were withdrawn from their cases and finally a few flaps of the abdomen freed the larva completely. There being no telson or anal thread this time, the larva at once left its old euticle and climbed upon the pleopod of the mother. The increased strength and size of the larva with the perfection of its limbs and candal-fan would make the danger of being lost on falling away from the egg much less than it was in the preceding stages THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 57 as the larva was now well able to follow its strong instinct to crawl upward and so recover its position on the mother, or, failing that, to live independ- ently, as it soon did. In the following description of the third larva we will emphasize its man- ner of association with the parent. In general form the young crayfish in this third stage (figs. 90, 91) is now for the first time like the adult, vet the great size of the eyes and of the ex- opodite seales of the antenne together with the very wide expanse of the tail fan and its long swimming plumes give the larva the aspect of a pelagic larva or of the adult of some lower form of marine decapod. In fact the ability to swim freely in great leaps which the third larva soon exercises and the trans- lucency of the body together with the big eyes and long antenne and very slender long legs and chele make the larva seem much like a small shrimp. The rostrum is now directed straight forward and is armed with large lateral spines that exaggerate the gothic style of the adult. All the appendages are now provided with long sete which make them sparsely hirsute as seen under the microscope, but only on the exopodite scale of the antenna and upon the telson and sixth pleopods are the sete set in rows that suggest a locomotor function. The use of the row of plumes on the seale of the antenna is not obvious but the long plumes all along the edge of the telson fan seem to add to its area effective in locomotion. The animal was still translucent and within it the liver showed as a narrow green band external to the large mandibular muscle, extending forward a short distanee and backward nearly to meet its fellow in the region of the heart. The sixth pleopods were now no longer inside the telson and the telson had become specialized as in the adult into an anterior and posterior part separated by a distinet movable hinge. Through the clear dorsal exoskeleton was seen the anus and a large mass of muscle on each side in the anterior region and in the posterior part of the telson the old radiating elands, which had eventually perfected the long sete that a high power showed to be plumose. The pigment cells represented by dark dots in figures 90 and 91, had be- come more numerous than before but were still chiefly arborescent red cells, though in many regions there were large blue cells not represented in the above figures though they now altered the ground tone of the animals. Posterior to the eyes the crowded pigment cells formed a streak along the base of the lateral spines and more to the rear a rounded area over the attachment of the great mandibular muscle. This round area on each side was especially conspicuous from the dense crowding there of blue pigment mixed with red. The dark ageregation of pigment cells avross the anterior edge of the abdomen still re- mained. The yolk area and its color were quite gone and on each side of the stomach a blue area indicated the gastrolith. Owing to the fineness of the 5 08 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS network of pigment cells on the tips of the chele these areas appear conspic- uously dark red. While the young in the third stage still remained on the mother for a while, they were free to move about and soon became more and more independent. A few hours after moulting some of the young were found walking about on the bottom of the aquarium. When disturbed they leaped backward and upward several inches by strong strokes of the abdomen and looked not unlike shrimp, while, when they walked, their habit of holding the anterior part of the body high up in the water as well as the attitude of the slender chele gave them the alert, nervous look of shrimp-like creatures and made them look very unlike the adult. The young climbed about on the pleopods of the mother, over her side and back, upon her mouth parts and eyes and, though at first densely crowded under the abdomen of the mother, more and more frequently they walked off to ereater distances, always returning to the abdomen if possible. The same larve that at times left the mother in the water, clung fast to her when she was lifted out of the water. But by violently shaking the mother in the water all the young could be shaken off. When two days after moulting into the third stage some sixty larve were thus shaken off and left in the same dish with her all but twelve had returned to the mother’s pleopods in twenty minutes. As many as a hundred young were seen walking about the aquarium and climbing up onto water plants for a time and later all but a few had returned to the mother. At first, however, the young did not wander far from the mother and when in a darkened aquarium the mother stayed in one spot for two hours the small fecal casts of the young over the bottom of the aquarium were almost all very near to the mother. In about a week this association of mother and offspring was gradually given up and more and more of the young failed to return to the mother. In nature it is possible that the mother and young separate quite soon, especially if the mother wanders about, but as yet nothing is known of the natural life of this crayfish when carrying the young. It may be that the female then lives in holes or cavities and then the young might long remain with her. In captivity the females with eges will dig holes in the mud and live in them as long as the water is well aerated, but come out when the water is not running. When kept in a small dish the female had no chance to escape from the young and in some cases some of the young remained with the mother and crawled over her three weeks and three days after leaving the egg, but when such females were put into a large tank they walked away and hid them- selves, leaving the young scattered about. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 59 Possibly the restricted artificial conditions both prolonged the duration of association of mother and young and lessened the variety of external stimuli so that there was a certain amount of domestication. By close confinement and by supply of abundant food to be given the mother and young without effort on their part, it would appear possible to make of the crayfish a domes- ticated animal with a more prolonged association of mother and offspring. Such experiments would build upon the ground that the crayfish is preéminently a creature in which embryonic life has been continued on as a series of stages, ancestrally free, but now dependent upon the mother and with special organs to ensure that dependence. The young that came down from the pleopods took food and eagerly de- youred fragments let loose from the mouth parts of the mother when she was feeding. At such times the young climbing on the mouth parts of the mother seemed in danger from those rapidly vibrating appendages but always seemed to be shoved aside and not devoured. When a couple of tubifex were put into the water and seized by a female, the young also took hold and either car- ried off pieces or continued to hold on while the worm was being dragged into the mouth of the female. Thus the young were drawn up with the food to the mouth of the female, but when between the maxillipeds the young leaped away and none seemed to be injured. The accompanying illustrations (figs. 92 and 93), being from photographs of living crayfish in water but poorly represent the pinkish mass of active larve crowding under the abdomen of the mother or the separate young elimb- ing over the back of the mother or walking about in the dish. They serve, how- ever, to show the crouching attitude of the mother, the size of the young in this third stage and the general character of this active dependence of young upon mother which was long since admired by Roesel von Rosenhof (1775), and might be likened to the clustering of hen and chicks or of sow and pigs. Roesel von Rosenhof’s statements regarding the association of the young and parent As- tacus are as follows: ‘Wenn die Mutter dieser kleinen Krebse, nachdem selbige sich zu bewegen angefangen, zuweilen bei ihrem Futter stille, oder sonst ruhig sitzet, so be- geben sich solehe von ihr etwas weg und Kriechen um zie herum; merken sie aber nur im geringsten etwas feindliches, oder sonst eine ungewonliche Bewe- gung im Wasser, so scheinet es, als ob sie die Mutter, sich zuruck zu begeben, durch ein Zeichen erinnerte; indem sie allezusammen geschwind unter den Sehwanz zuruck fahren, und sich wieder auf einen Klumpen zusamensetzen, worauf sich die Mutter sammt selbigen mit mogligster Hilfortigkeit, in Sicher- heit begiebt, welche sie aber etliche Tage darauf, nach und nach verlassen.”’ The suggestion of a possible signal to recall the young raises the question as to the nature of the means by which the young associate with the parent. 60 THE -YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS The interrelation of mother and young in the crayfish would seem a profit- able field for study of comparative family life and the following few facts sug- gest lines of inquiry. It is not evident that the mother gets any advantage from the clinging of the young to her body, though if in the third stage they eat off the old egg stalks and eases, the cleansing of the pleopods so caused might satisfy in the mother something akin to whatever it may be that causes her to laboriously cleanse that region just before the eggs are laid; but judging from the long endurance of the load of eggs and larve and from the frequency of dirt and parasites upon the pleopods it seems hardly probable that the female would feel an in- stinet gratified when the pleopods are clean again. That the female is at all conscious of the presence of the larve remains to be found out by evidence not yet at hand; however, when a lot of young were put into a dish with a male and they climbed over him he cleaned them off from his abdomen and seemed to be annoyed by their climbing over his head, but when the same young were put with a large female that had reared her own young long enough and natur- ally separated from them some weeks before, they climbed up onto her without her showing any evident sign of being affected by their presence upon her chelx, thorax and abdomen. When a female was feeding and the young climbed upon her chele she did not seem aware of them but reacted eagerly to a tubifex near her chele. Whether, then, the young are simply tolerated, [as are the small parasitic leeches over the body of crayfish,] as a continuation of the eggs that are part of the body, or whether the females have special responses to stimuli given by the young is not known, but the former seems not improb- able. The young, on the other hand, act as if they were strongly affected by the mother but this may be due to a few simple reflexes and without any complex visual or other conception of the mother’s existence. At all events there is no evidence that the young distinguish one female from another and when two mothers were in the same dish some young of one climbed up onto the other. Even when the young shaken off a female when they were in the third stage were put in a dish with a female carrying her young in the second stage they climbed up amongst the younger larve of the strange female and seemed con- tent. When the young were shaken off two females in separate dishes and the females exchanged, the young climbed up onto the strange females ; in a few min- utes all but three were upon one female and all but seven or eight upon the other though she was moving about. Some young C. affinis in the second stage were taken off from the mother and put with some young of C. Diogenes of the sec- ond stage also removed from the mother and all were then put into a dish with another female C. Diogenes hearing young in the second stage. All the young got upon the foster mother and continued there, though the C. affinis were THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 61 searcely more than one-half the length of the other larve. The young that are shaken off dart, swimming, about in the water but soon settle to the bottom and climb up upon one another in heaps or try to climb up at the corners of the dish and upon water plants. The tendency to climb onto the mother is very strong so that when a female walks alone swinging her pleopods laden with young she often gathers up free young that are walking about, as these when touched by a pleopod may not spring away but turn and climb onto the mass. In fact early in the third stage a larva that fell to the bottom, apparently for the first time, stood with head end elevated and quickly responded to the presence of a passing pleopod. Even if lying upon their backs they quickly seize and mount the pleo- pod of a passing female. A piece of white cheese-cloth, however, did not appear to stimulate them to climb upon it and when a lump of rough cement, a model of a toad, was left in the aquarium very few got on it, most of them preferring the mother. Possibly the young receive chemical stimulus from the mother that aids them in returning to her. That they responded to some chemical stimuli is most probable and the following facts may be interpreted on this basis. When the mother feeds, the young gather under her mouth and even after the food is gone they remain as if excited by chemical substances coming from the mouth parts of the female. It was also found that while the young climb onto a dead female and even over the exposed surface of the freshly broken abdo- men, after some days they remain on the rest of the surface but not upon the broken surface where chemical substances due to decay probably existed. During the time the larve in the third stage associate with the parent the large collections of old egg shells and skins on the pleopods disappear and are probably eaten up by the larve since their intestines contain minute sete that might come from their old cuticle and since in Astacus, Chantran (771) found that the larve eat their shells and cast off cuticles. It is therefore possible that one reason the young remain with the mother is that this food supply on the abdomen acts as a stimulus to them. However, along with whatever chemotactic movements there may be, there are other factors concerned in the association of the young with the female as indicated by the few following experiments, which tend to show that the young are controlled in part by responses to light and to gravitation and con- tact. Thus when a small paste-board table was put under the water the young collected on the under side of the horizontal paste-board and remained stand- ing upon it, in an inverted position. When the paste-board was then turned up- side down the young scattered in a few minutes so that half were off the paste- board and some were under it in its new position. In crawling under such a table the young reared up and seemed to be trying to reach it above their heads as if 62 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS influenced by it before they touched it. When a small glass table was used the young also collected under it and reached up as if trying to climb up to it. When dark paste-board was laid on part of the glass, above, twenty-two larve had collected under that part of the table shaded by the paste-board, in two hours, and none under the clear part. It may be imagined from these indications that the larve in the third stage have a tendency to climb upward due to negative geotactic responses, that they prefer the shade as being photophobie and that they come to rest standing upon a solid object as being thigmotactic. Assuming some such responses together with some chemotactic phenomena, possibly all the habits of the young in refer- ence to the mother may be explained; possibly not. The association of the young and the mother, however brought about, endures in Cambarus affinis during two larval stages and a part of a third. In Astacus leniusculus, however, it is the second and not the third stage which gives up the association with the mother. Whether this is a generic difference cannot be determined without extensive observations, as almost no data are yet at hand. That Cambarus in general have three dependent stages seems prob- able from the following few facts that have been recorded. Faxon (’85) saw specimens of C. Clarkii 7 mm. long upon the abdomen of the mother (as previously recorded by Hagen, ’70), in which the tail-fan was perfect so that with the ability to lead a free existence there was still the habit of associating with the mother. In C. affinis larve in the first stage are 4 mm., in the second 5 mm., and in the third 7-8 mm. long, and it seems probable that the above recorded C. Clarkii were also in a third stage. The same author saw upon the mother C. gracilis 9 mm. and C. Bartonii 10 mm. long, with perfect tail fan. These two species may well have three stages of association as in C. affinis. Also he records C. rusticus just hatched and 4 mm. long as being so embryonic that we may assume they would remain with the mother as long as C. affinis does. On the other hand Steele (:02), who first observed the living larve of Cambarus associated with the mother, records C. gracilis 7 to 8 mm. long with a bent rostrum as if in the first stage though so large. And in a more detailed study of C. virilis she deseribes the first stage and then a second stage nine days oid with the tail fan complete and other features of the third stage of C. affinis, but as she says the larva six days old could swim it seems prob- able that in both these species as in C. affinis there are three stages in the period of association, the second of which is easily overlooked. There is then no obstacle to assuming that we may expect to find in Cam- barus that the larve remain with the mother through a first, a second and part of a third larval stage and that in Astacus they remain through the first and part of the second only. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 63 The active third larve becoming independent lived some eighteen days in all before moulting again to pass into a fourth stage and these in turn were followed through more stages till eventually adults were reared in the labor- atory. Without describing the gradual changes of the early larve to the details of adult form we may now give a more complete record of the rates of growth of young crayfish than has hitherto been possible as a number of new observa- tions have been added to those previously published, Andrews (:04), and certain points as to the period of sexual maturity determined. We will give certain facts as to the number of moults, as to the rate of growth, as to the develop- ment of the external male organs, as to the ratio of the sexes and as to the ar- rival of sexual maturity and the ability to breed in captivity. Observations made upon the young of a single female which laid March 28, 1903, and whose eggs hatched May 18, 1903, gave the times of moulting and rates of growth of the larve represented in the following table: Stages. Dates. Duration.| Size. | Increase. Habit. Ist stage. | May 18-May 20; 2 days | Alia || o 6 oc Attached to mother. 2d stage.| May 20-May 26| 6 days 5 mm. 1mm. | Attached to mother. | | \ Associated with mother for 3d stage. | May 26-June 15) 18 days 8 mm. | 3mm. about one week and then free. 4th stage. | June13—July 1) 17 days 12mm.| 4mm.| Free. 5th stage. | July 1-July 6) 5 days | 15-18mm.)} 3-6mm. | Free. 6th stage. | July 6-July 17) 11 days 21 mm.| 3-6 mm. | Free. 7th stage. | July17- ? 2 29 mm. 8mm. | Free. | These same young were kept till October 6, 1905, without observation and the eight survivors then measured as follows: 62, 55, 53, 49, 50, 45, 43, 41 mm. in length. The average is nearly 50 and as far as can be judged from the above table such a large larva as that measuring 29 mm. in the seventh stage would probably have passed through at least four or five moults to become the large one 62 mm. long and smaller larve would have required as many or more moults if the increment was as large as in passing from the sixth to the seventh stage; but if the increase was the average for the above seven stages, searcely 4 mm., then at least six more moults may have taken place. Other data show that the rate of growth is very different in individuals and in some cases seemed to depend directly upon food supply so that the number of moults during the first summer is probably not by any means constant but different in individuals. However, we are sure that at least seven stages may be passed through and probably eleven to thirteen larval stages may occur in the first sum- mer of the creature’s life outside the egg. Yet data given below show that the larvee may remain but 20 mm. long in the fall, owing, probably, to insufficient food arresting them when in the sixth stage. 64 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS After October there were no moults till the following warm season, the lengths remaining constant during the winter. Just as the growth in the first summer was restricted to the five months of May, June, July, August, and Sep- tember, so in the second summer there was an increase of 25 mm. during that time and though the moults were not observed there may well have been four yr five during this second summer. In a few exceptional cases a moult took place in the early spring before May and a gain of 6 mm. was observed then. After a second winter of no moults there followed a third summer of growth in which a single observation showed that a specimen 70 mm. long December, 1902, was 76 mm. long July 1, 1903, an increase of 6 mm. probably due to one moult. As this same individual increased to 90 mm. by October, 1903, it prob- ably moulted two or three times or in all three or four times during this third summer. We thus have evidence for seven and probably as many as twelve moults the first summer, four or five in the second summer, three or four in the third summer, and in the ues summer perhaps only one or two moults to judge from a single case of 8 mm. increase. Without reference to the number of moults, observations upon the amount of growth of young crayfish hatched from eges laid in the laboratory and kept under various artificial conditions in the laboratory, were made and are re- corded below as a substitute for data never yet obtained as to the rate of growth of American crayfish in the open. é The following list gives the length in millimeters from tip of rostrum to end of telson of 101 young hatched in May, kept in six different tanks in different years and under somewhat different conditions and measured in October when four months old. I. 62, 58, 53, 49, 50, 45, 43, 41 MG 60) 5952702 439 40a aoe adel Ti, 41, 38, 24. 56. 32; 44, 43, 39° 39) 33, 35, 36, 32; 34, 30) 23.130) 29) 27, 32: IV. 47, 39, 33, ce 20), 28, 31938..a0. SIN ose On Soro amonerds us 1, 46, 52, 46, 45, 46, 41, 39, 40, 3¢ V. 48, 51, 41, a 43, 37, 38. VI. 55, 48, 55, 48, 38, 50, 40, 40, 53, 46, 40, 42, 40, 43, 50, 45, 44, 48, 50, 49, 37, 57, 44, 45, 43° 46, 46, 36, 38, 38: It is obvious that there was great individual difference in size attained in the first four months of life both under different conditions and when kept in apparently the same conditions, but of course subject to different chances of food supply even in one tank. The first group of eight survivors ranged from 41 to 62, with an average of nearly 50. The second group of 10 survivors ranged from 22 to 62 with an aver- age of 43. THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 65 The third group of 20 ranged from 23 to 56 with an average of 35. The fourth group of 26 ranged from 20 to 47 with an average of 36. The fifth group of only 7 ranged from 37 to 51 with an average of 43. The sixth group of 30 ranged from 36 to 57 with an average of 45. The large size of the crayfish in the first group was undoubtedly due to greater care taken to feed them and the differences in other groups were prob- ably due to differences in food supply and also in temperature. Judging from the above 101 measurements the young four months old may vary in length from 20 to 62 mm. but while the average of all was 41 mm. the average of different experiments ranged from 59 to 50. We may then expect this crayfish to be nearly two inches long in the autumn of its first year of life. Measurements made in the autumn and again in the next spring showed that there was no growth from October to May, except in rare cases in which a moult occurred in early spring. The crayfish thus started in their second summer of life with a leneth of nearly two inches and were the same size when twelve months old as when four months old. The rate of growth in this second summer may be inferred from some few measurements made in October upon crayfish reared from eggs laid in the laboratory and measured then, when sixteen months old. Of five larve measur- ing May 26, 1904, 62, 55, 53, 50, 62 mm. the three survivors in October, 1904, measured 75, 80, and 72 mm., an average of 76 mm. From the average, 58, of the above five to the average 76 of the three survivors there was an increase of 18 mm., or a gain of 30 per cent in length in this second summer. And records showed that one of these larve had grown from 62 to either 75 or to 82, that is added 13 or else 20 mm. to its length, or increased 21 per cent or else 32 per cent. Another group of young left three survivors of 56, 70, 75 mm., or on the average 67 mm. when 16 months old to represent the seven which were 40-45 mm. long when twelve months old; they had thus probably added 25 mm. or gained more than 50 per cent in the second summer. A third group of young, twenty in number, ranging from 23 to 56 mm. when twelve months old, left only two survivors at sixteen months, which measured 70 and 79 mm., or an average of 74: there was thus an increase from the average 35, of 39 mm. But records of the individuals showed that the few survivors were probably 43 and 56 mm. long in the spring and had grown only 93 and 27 mm. in each case which would be an increase of 53 per cent and of 63 per cent. The above few data indicate a growth in the second summer of sometimes 50 per cent of the length; that is a erayfish two inches long the end of the first summer may be expected to be three inches long the end of the second summer, 66 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS when sixteen months old. But here again the individual differences are very ereat. In the second winter there was probably no growth; two crayfish 75 and 80 mm. long in October, 1904, were the same length in May, 1905, and, in gen- eral, moulting during the early spring was rarely observed. The only observation upon the growth of crayfish reared from the egg and kept during the third summer was the following: two individuals 70 and 79 mm. long when twenty-four months old left one survivor 90 mm. long in October, 1903, when 28 months old. It had thus increased either 11 or else 20 mm. in the third summer, that is, it added only 14 per cent or 20 per cent to its length. Probably then a crayfish may grow in the third summer from a length of three inches to be somewhat short of four inches. The one young crayfish kept during its third winter did not change but measured 90 mm. in October, 1903, and the same again in May, 1904. In its fourth summer this crayfish grew to a length of 98 mm. by October but died then when three years and four months old, having grown but 8 mm. or barely 9 per cent in this summer. This may indicate a great diminution in rapidity of growth as the maximum size is approached so that the crayfish one hundred and twenty millimeters long that are taken in the Potomae may well be six or seven years old; but from one specimen we cannot say that the rate is naturally lessened in the fourth year and a possible yearly inerement of one inch would make the large crayfish of five inches only four years old. The age of Cambarus affinis would seem to be roughly determinable from the formula A = L— V, where A= the age in years ending in May, L= the length in inches and V = one, in conditions of maximum favorableness, but in unfavorable conditions V may become zero and in very old large crayfish V may be a negative quantity. Soubeiran (765) gave the length of Astacus in France as 50 mm., 70 mm., 90 mm., 110 mm., 125 mm., in the first to fifth years inclusive, that is with an annual increment of 20 mm., which, however, became less in the fifth year, so that he could not tell the age of crayfish 160 mm. long nor of any very old ones 190 mm. lone. Compared with this Astacus, C. affinis starting as a much smaller egg caught up in size in the first summer and for two years showed just about that same increment of 20 mm. annually, but then probably began to grow with more diminished speed than did the French Astaeus. The sexes of these young crayfish are early determined and in the third larval stage the female shows the beginning of the annulus and the male the external openings of the deferent ducts, as elsewhere described (Andrews, :06). But the difference between the appendages of the first abdominal somite in THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS 67 male and in female are first well seen in the fourth stage when the female still bears minute tubercles and the male long papille to represent the appendages of this somite. From that time on to the autumn the distinctive characters of the sexes become perfected so that the annulus of the female is much like that of the old erayfish, and the first abdominal appendages, or stylets, of the male are also much like those of the full grown erayfish. It is thus very easy to recog- nize the sexes by external characters when they are four months old. The actual length in millimeters of the male stylets in crayfish of four months is given in the following table: Length of body . . . 55 53 49 4 Length of Ist stylet . 9 9 9 Length of 2d stylet . 10 10 9 43 60 52 52 4 On 5 39 40 33 30 7 6 6 5 4 if Af © aay a! CO ~l a oS 5 wo} 1 LO SLOSS =~] The second stylets are thus longer than the first and as much as 18 per cent to 20 per cent of the length of the body so that the stylets are in about the proportions they will be in larger crayfish since specimens 115 mm. long have stylets 18 and 22 mm. long. Some examinations of various catches of adult crayfish indicated that there is no great disparity in number of males and females and though it is not known whether one sex or the other is subject to greater mortality in early larval life, the following observations tend to show that the sexes are about equal when four months old so that it may be that the eggs are about equally male and female. Observed at four months: of 26, there were 15 male and 11 female; of thirty, 11 were male and 19 female; of 7, 4 were male and 3 female; of 19, 10 male and 9 female; of 10, 8 male and 2 female; of 8, 5 male and 3 female. Of the entire 100, 53 were male and 47 female. With no marked disparity in numbers and with well formed external organs the two sexes when four months old and about two inches long often have well developed sexual instincts. Thus in October a male 55 mm. long, 15 mm. wide, with tail fan 22 mm. wide, antenne 48 and 51 mm. long and entire expanse from tip of chelx to end of telson only 75 mm., was seen to try to con- jugate with a sister 62 mm. long when four months eighteen days old. An- other female had a mass of sperm transferred to its annulus by a male of like age when four months old and 57 mm. long. Many cases were seen in which the four months young had conjugated, thus of 19 females four months old, in one aquarium, five bore sperm given by males of like age. Though the sperm transferred by these young males was apparently perfect it seemed doubtful if these unions would lead to fertile eggs but when a number of four months females thus provided with sperm by males of like age were kept isolated from all males during the winter they laid eggs in the spring which developed and thus demonstrated not only that the females could lay eggs when but a year 68 THE YOUNG OF THE CRAYFISHES ASTACUS AND CAMBARUS old but also that these eggs would develop after union with males of the same age, union which moreover took place in the previous autumn so that these ex- periments also showed that the sperm got in the autumn and kept all winter would fertilize the eggs, parthenogenesis being, supposedly, out of the question. In Astacus, Chantran stated that the eggs were laid only ten to forty-five days after union but in Cambarus affinis as many as 230 days may elapse. The data on which the conclusions rest are as follows. ee : seul 2 PY Se ie i : _— 5 oe ; 1%. ty = 7 t 7 _ ‘= os 7 » a 8 =e é . - “a 4 . ° @ as ~ «6 — - al 7 =e | > i - = ’ Aw ‘ oe —_ ve 7 4 x r . on « » algphs he , 7 : - ro f “s _ 7 : —— ll 7 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE—ANDREWS PLATE V ~< ASTACUS LENIUSCULUS: SECOND STAGE FE. A. AnpreEws, del. SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE—ANDREWS PLATE VI ~ Se : oo SSSS= Rs = =: = SESS ——— aed SST [OE are cea =< = : SS — a ASTACUS LENIUSCULUS: THIRD STAGE E. A. Anprews, del. SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE—ANDREWS a ee 2 PLATE VII CAMBARUS AFFINIS: FIRST STAGE E. A. Anprews, del. SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE—ANDREWS PLATE VIII 61 CAMBARUS AFFINIS: FIRST AND SECOND STAGES E. A. AnprREws, del. SMiTHSONIAN CONTRIBUTIONS TO KNOWLEDGE~ ANDREWS PLATE Ix CAMBARUS AFFINIS: SECOND AND THIRD STAGES E. A. Anprews, del. sow , ried eee Pee SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE— ANDREWS PLATE X FIGURE 93 CAMBARUS AFFINIS: THIRD STAGE E. A. AnpDREWS, photo. : SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE ; PART OF VOLUME XXXV__ A MONOGRAPH OF THE — SYNAPTIDA AND MOLPADIIDA _ Including a Report on the Representatives of these Families in the nk Collections of the United States National Museum BY phi: HUBERT LYMAN CLARK BEBE INCR Ey. 200000002 (No. 1723) CITY OF WASHINGTON vey PUBLISHED BY THE SMITHSONIAN INSTITUTION ‘ 1907 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE PART OF VOLUME XXXV tie ODOUS HOLOTHURIANS A MONOGRAPH OF THE SYNAPTIDA) AND MOLPADIID Including a Report on the Representatives of these Families in the Collections of the United States National Museum BY HUBERT LYMAN CLARK CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1907 Commission to whom this memoir has been referred : ADDISON EMORY VERRILL WILLIAM KEITH BROOKS WASHINGTON, D. C. PRESS OF JUDD & DETWEILER, INC. 1907 ADVERTISEMENT. The present memoir by Dr. Hubert Lyman Clark, of the Museum of Comparative Zoélogy, in Cambridge, Massachusetts, on ‘‘The Apodous Holo- thurians,’’ sea-cucumbers or trepangs, forms part of Volume XXXV of the Smithsonian Contributions to Knowledge. In this memoir the attempt has been made to give a complete summary of our present knowledge of the two families of sea-cucumbers which lack tube- feet. As a system of classification is the essential foundation upon which any discussion of structure, habits or relationships must be built, the first section is devoted to a discussion of the history of the classification of the two families (Synaptide and Molpadiide), with comments on the principles involved and a final summary of the system adopted. Most of the original investigations hay- ing been based on material in the collections of the United States National Museum, the second section is an annotated catalogue of the Apodous Holo- thurians of that institution. The third section deals with the family Synap- tide, and takes up in regular sequence, so far as our present knowledge per- mits, the structure, physiology, development, habits and classification of these animals. In the last subdivision of this section, each species, recognized as valid, is treated separately in some detail. In the fourth section, the family Molpadiide is treated in the same way. The most important feature of the work is the recognition of the changes taking place in the maturing and senescence of individual holothurians, par- ticularly in the family Molpadiide. As a result of this, radical changes im nomenclature have been necessary, but every effort has been made to have the system adopted accord with the most widely accepted codes, and thus be as stable as possible. Special attention has been given to geographical distri- bution, but the work in this line is chiefly of value as a summary of our present very inadequate knowledge. Artificial keys to genera and species have been freely used with the intention of making the work as useful as possible to all subsequent investigators, and the numerous figures, most of which are copied from other writers, are given with the same end in view. {nm accordance with the rule adopted by the Institution, the work has been submitted to a commission consisting of Prof. A. E. Verrill, of Yale University, and Prof. W. K. Brooks, of the Johns Hopkins University, who recommended its publication in the present series. Cuas. D. Watcorr, Secretary. SMITHSONIAN INSTITUTION, Wasuineton, June, 1907. 3 TABLE OF CONTENTS. liminaalniiont ch oo oko ade ob 6 oebe Od 0 Onn neitrOl cine SOmOdc > Oo omem tod GUC OU GDC pOnadc Parr I. The Classification of the Apodous Holothurians ............-..++0+ +e eee ete Table of Accepted Genera, with Type Species .......--.- +2 e sere eee cette reece Part II. The Apodous Holothurians of the United States National Museum......-..+++--++-++s Sommmiliidlis vce cobs sonoba rode Ok Come ns an nHene OOO Ong eo Die DMD DUD Mang. MIGIRG HIGHS 3 eAee nto cn eddugd Gc a8 cae Conan obo eCard Oren Dan DCU uaaicD GoD Wie slnOlaasy gg oe oe koeso5 100 G0 GOO On Bat OOD an A eInEn at cin De cataract aps iRlonian aimél SivO.o oc cdsnccogeoandnengancoosocmogonoceo PAusEe Go DeehA DOGO OCE Clim soaccococdansecduacod0 pobOgduoceg heen o ano UpUE COO mb a soo COU CUE Oc,0 OCI [BOG SU EIGS 6 Sao nas oo noo eeE pp ae eri tGaln Ure oer as OC Gace TMAMGIES 4.0056 coon wo en CRAB OOO EIOO IRIE O & Gorin Rite n nae a eoro. cee nus ama ORCC SWURE-ORECING, 56 cio bon H Leto Uene IO OD Oro Cee Dean Oi iC SO niece Cn a cca LAcIIROIHECH? 45h go coneeEee age neee a aan Op aEeoc oor eerie Pasiitiaell ....ccac0ce nan son nso od GNds oAbmood quo Soe apaOnUOUU COD GUN OOUCUIC Guigitilos? soe.c0k ae 4a adore Uemieerceo samira an ee Ten arene adr ac 05.0, 0.0 TGS oacceceeas lOesod Oa er ama a ae OlmcDIOinne ee OOncs ci OU COCO accion NIGETOME SWISH nee /6 6 dose bio ele ge eae ie oReeion SRI Omit aC Boakeralll, chee occ son ete ene On ORs Energi ei ero Cc a Sa [SERENE cee cs ch necod Oe Oe en noNG ne ab DOC e rigor econ Ona aa @onnechivetssucl Mayer aoc s 32s o ee set ee eae eens enneonome ses TRignaimeallls , oeden cHege dee GaceGr 40 Sep oo ood cond OUD UU UOC MEE Callemeat) 10a (asi) ep pee @an0be ae anne eee Oreo nop COUSoo SOR OOIGo OG aging maGl WW soceucenesunaaecssouc59ec0005 DUD DOU OC odDOOUD Wiha) cb doonnowwe doc ceo su oeomeeedee cadomaccuUrGdcD ac cInO UG TOC Ciamradl Reds ceccuvocsscccoodooesmccdspoauoogoDodddoccDOSOgBdC MurilnepryaGrantllesis tec ce 2 eco 2 erie rns hoe oni Climeuilane IMGREIES cé.doc006 cco ooame ono nnoneGacouddodooDU Ud DUdBO ODO CCOGCCICC ionpibudinal Muscless.\. 12... 2-2 e-+ oe cme ee esate acne deste meee ct cee oe Tigingy TDyeiWAGIIGi 6 Ga egctaee eae neo OA OOG. Seine I ter aa ee ca acaiaieaac TBQdieGbTUy a pocde donb O60 tie Coron Crmneria oy cack gnc Cen antic Oca vei Spllemanne RING . cnc ogee noe Oe Cee ania ta anc Daer astra aa ied ce Chinliacinatis, TMs Gc 6 dabeba od ed or annedG re nora e Gao sD G Uno onD aia aacie Winicenagenllr Si RuGi, 9 oanee > boo aes ERERI ie eo UIC ia Raa ara limgniiney (CHGaiL “ype Goece 55 tnnCe Oe eee Oi Ot a aCe Rta i tar iano Diane nae Cllkaiedl Pimms ccoéccancocepoeue ao onasgeod Coded oO GUC odroU OU DOO GRD CD UOT TSlnad, SWRETI, oho BA ceo eee Se Dre IOI renee eee CIE A Sac ica atc TPejonGricuiye: SyG init ee Oe Cha ore SIC Ta a ceca ca a Bpnibrallanye cs crn be oie OSES IB S1Oc0 Ch ne ICI ne ND ca aaa Page ili 22 23 23 dl 42 42 42 45 44. 46 46 46 46 AT 47 48 6 THE APODOUS HOLOTHURIANS Page Physiology asians satis oats ces acres. eles peebne G SR oon ORCA nee CEO Cees 61 Motion « cckcs ion ele tars eicves Slonshe eiinelanic bene EIR eo EO OOS See Oe Cert 61 Digestion and! Absorption. sc. so sesinete Gate a ine cheers eee err 63 Circulation and ONiibritlowis-.: iy. se crsre sees eens ie fae Sele eee oe ee 63 Respirabloms $24 fer 33. cls.c See sones,e hers e e nr ee Pe cae ae OEE eer 64 1D 1) ee eae Ince Ae Orco Oa tcomce ad noom God faces nGomacdads cand 64 MO CMSAELOM va, irs ausyensis sts auever sue ove lere ic hel GxStLe) pe em RereTe Eo e Rone Sn ea 64. Regeneration «soso. wine at crncte wiste.coates om ously e olewacrceeicetie ic os Conon ene Ree eee: 65 Reproduction slya cr weserenva iiss sists oer terenst slepe oe a ICC eee 66 COO DY. ais forest blo tore tapeuoen ats Sievione ase foueuanee oTovae eevee arte RCE Ter RoI eno ne ea 66 10) ko) Eee oe eee we a ae ANA aNcieA esd ob Soa bucon coded Sc 68 Iey, tosthe}S ubramlliess ots Siyniap tid ce sree eee cree iene ee ete 70 Sh nel o] nhl ae aa MOC ROAD Oma One moe com Y Goma cratapednnpacas 70 Mraapta oe.2 3:2loce 52sec sioiuedels oper ae eee ae ea ene eee 72 Opheodesoma: sis ecco cas Ateises oer a eaees ogee SO Ee ee eee 7: Polyplectamar [ei 8, Wie, Gh Sasi, UMA I Antero-posterior diameter of bone, 2 mm.; lateral, 2 mm. Antero-posterior diameter of medullary canal, 1.4 mm.; lateral, 1.5 mm. The medullary canal is full. Medullary index, 111%. Structure.—The section shows the three divisions, external circumferential lamelle very thin, central ring of lamella, and internal circumferential lamelle. The external lamelle are little more than a condensation of the external portion of the central ring. The central ring consists of concentric lamelle with oval lacune. In the posterior and part of the lateral walls, oval lacune with bushy canaliculi are crowded together between the central ring and internal lamelle. A few canals appear in the inner wall. The internal circumferential lamelle form a narrow ring around the medullary canal. Type I-III, Ia. RIGHT FEMUR OF BUFO LENTIGINOSUS WOODHOUSH. NO. E 40 9, U. S. NAT. MUS. IPTao) HIGs Sones iNeRLABes! Antero-posterior diameter of bone, 2.5 mm.; lateral, 2 mm. Antero-posterior diameter of medullary canal, 1.5 mm.; lateral, 1 mm. The medullary canal is full. Medullary index, 45%. Structure.—The section is composed of a thick ring of concentric lamelle with oval lacune, crossed by numerous radiating canals and perforated by a few Haversian canals (Ia). Internal circumferential lamellae form a narrow ring around the medullary canal. Twofold division. Type I-III, Ia. RIGHT FEMUR OF BUFO AMERICANUS. AMERICAN TOAD. AMER. MUS. NAT. HIST. Pu. 3; Pia. 36. Syn. Tas. 1 Antero-posterior diameter of bone, 2.5 mm.; lateral, 2 mm. Antero-posterior diameter of medullary canal, 1 mm.; lateral, 0.6 mm. The medullary canal is full. Medullary index, 15%. Structure.—The section is surrounded by a wide band of lamelle with oval lacune and bushy canaliculi, interrupted by frequent Haversian canals of the (la) differentiation. The band is widest in the anterior and imer walls. No. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE 47 Underneath this band is a single lamina with long, well-developed lacune and straight canaliculi, parallel with neither surface of the bone, but occupying an irregular position. Underneath the lamina is a wide central ring of lamelle with oval lacune and bushy canaliculi, interrupted by frequent Haversian canals of the (la) differentiation. A narrow ring of internal circumferential lamellae surrounds the medullary canal. Threefold division. Type I-II-III, Ia. RIGHT FEMUR OF BUFO LENTIGINOSUS CoGNATUS. No. 13 11, U. Ss. NAT. MUS. Binoy WiG.ce (ae nowNie Absa Antero-posterior diameter of bone, 2 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 1 mm.; lateral, 0.9 mm. The medullary canal is full. Medullary index, 42%. Structure.—The three divisions are evident. The external circumferential lamelle, with long lacune and straight canaliculi, surround the section. The central ring is composed of lamelle with oval lacune perforated by Haversian canals of the (Ia) differentiation. The internal circumferential lamelle form a narrow ring around the medullary canal. Type I-III, la. RIGHT FEMUR OF BUFO VALLICEPS. NO. E 21 5, U. S. NAT. MUS. Pt. 3, Fic. 38. Syn. Tas. I Antero-posterior diameter of bone, 2 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 0.9 mm.; lateral, 0.8 mm. The medullary canal is full. Medullary index, 317%. Structure.—The section is surrounded by a thick band of lamelle with oval lacune and bushy canaliculi. It is perforated by numerous Haversian canals of the (Ia) differentiation. In the inner wall many radiating canals appear. The canaliculi from adjacent lacune extend into both the cireular and radiating canals. A narrow ring of internai circumferential lamelle encloses the medul- lary canal. Twofold division. Type LIII, la. RIGHT FEMUR OF RANA BOYLI. NO. F 61 20, U. S. NAT. MUS. Pr. 3; Hie. 395 Syn. TAB] Antero-posterior diameter of bone, 1 mm.; lateral, 0.6 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.4 mm, The medullary canal is full. Medullary index, 47%. 48 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 39 Structure.—The section is composed of two parts, viz.: A wide ring of external lamellae with oval lacune and bushy canaliculi, and a narrow ring of internal lamelle with long, narrow lacune and straight canaliculi around the medullary canal. Twofold division. Type I. VIL. REPTILES Thirty-four femora were examined. GENERAL CHARACTER OF THE K'EMUR The general shape of the reptilian femur varies considerably. The tri- angular, elliptical, round, and indeterminate forms are present. The majority are elliptical. In some sections the antero-posterior diameters are longest, as in many of the lizards, and in others the lateral diameters are longest, as in the turtles. The contents of the medullary canals are variable in character. In the lizards they are filled with marrow and in the turtle with cancellous bone, the meshes of which are filled with marrow. The medullary surfaces are smooth in the small, and rough im the large, femora. The medullary index varies from 0 to 88% with an average of 26.1%. Excluding the turtles, in most of which the index is zero, the average is 33%. The reptilian femora follow about the same kind of development as was seen in the amphibians, but the development is carried further. The first type of bone predominates in both amphibians and reptiles. Ha- versian canals, (Ia) stage, appear in the amphibians, while a better developed form of Haversian system, (Ib) stage, is found in some of the reptiles. In the amphibians the external, internal circumferential lamelle, and central ring appear, while in the reptiles these divisions are not prominent. Cancellous bone was found in one amphibian, the Amblystoma. It does not appear in the lizards, and is ‘a characteristic structure of turtles. DeraILep EXAMINATION RIGHT FEMUR OF SPHENODON PUNCTATA (MOST PRIMITIVE OF REPTILES ). AMER. MUS. NAT. HIST. Pu. 3, Eig. 40. Syn. Tas: I] Antero-posterior diameter of bone, 38 mm.; lateral, 2.5 mm. Antero-posterior diameter of medullary canal, 1.5 mm.; lateral, 1 mm. The medullary canal is full. Medullary index, 26%. Structure.—The section is composed entirely of concentric lamelle with oval lacune and bushy canaliculi surrounding the medullary canal. A large No. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE 49 vascular canal appears in the inner and posterior wall. There is very little differentiation of structure. The bone is uniform. Type I. RIGHT FEMUR OF PHRYNOSOMA CORNUTUM. TEXAS HORNED TOAD. No. 1200, AMER. MUS. NAT. HIST. Pr. 3; Fie. lo Syn. Tas, 11 Antero-posterior diameter of bone, 2.8 mun.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 0.8 mm.; lateral, 0.5 mm. The medullary canal is full. Medullary index, 10%. Structure —The section is composed of concentric lamellae with long and oval lacune and straight and bushy canaliculi surrounding the medullary canal. The lacune of the external portion are long and narrow with straight canaliculi, and those of the medullary portion are oval with bushy canaliculi. There is very little differentiation of structure. The section shows an indistinct twofold division. Type I. LEFT FEMUR OF CHAMELEO VULGARIS. CHAMELEON. NO. 135, AMER. MUS. NAT. HIST. 1g, 6y Wbies GS Sivas MA IIL Antero-posterior diameter of bone, 1.8 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 1 mm.; lateral, 0.8 mm. The medullary canal is full. Medullary index, 42%. Structure.—The section is composed of an external ring of lamelle with oval lacune and bushy canaliculi. The lacune tend to flatten as they reach the external surface. In the posterior wall the lacune are large and nearly round. Underneath this ring is another of lamelle with oval lacune and bushy canali- culi. The two rings are separated by concentric, central lacune closely packed together. The bone shows an indistinct twofold division. Type L RIGHT FEMUR OF PHRYNOSOMA DOUGLASSIT. No. L 50 12, U. Ss. NAT. MUS. Pr. 3, Fie. 48. Syn. Tsp. IL Antero-posterior diameter of bone, 2 mm.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.5 mm. The medullary canal‘is full. Medullary index, 12%. Structure.—The section is composed of concentric lamellae with oval lacune and long canaliculi surrounding the medullary canal. Little or no differentia- tion has oceurred. The section has a uniform structure. Type I. DO SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VoL. 35 RIGHT FEMUR OF PTYCHOZOON HOMALOCEPHALUM—GECKO, NO. 684 AMER. MUS. NAT. HIST. Pi. 3. Bie. 445 sya CDAB. Antero-posterior diameter of bone, 1 mm.; lateral, 0.8 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.4 mm. The medullary canal is full. Medullary index, 33%. Structure —The section is composed of concentric lamellae with long lacunze and long, straight canaliculi surrounding the medullary canal. Very little dif- ferentiation appears. The section is uniform. Type I. RIGHT FEMUR OF IGUANA TUBERCULATA. AMER. MUS. NAT. HIST. IP it, Sh ihe, ala, Shani, UME HI Antero-posterior diameter of bone, 3.5 mm.; lateral, 3 mm. Antero-posterior diameter of medullary canal, 2 mm.; lateral, 2 mm. The medullary canal is full. Medullary index, 61%. Structure.—The section is composed of concentric lamelle with long lacune and long, straight canaliculi surrounding the medullary canal. The posterior wall is thickest and shows a column of oval lacune and their lamella extending from the medullary canal toward the external posterior surface. Very little differentiation of structure appears. The section is uniform. Type I. LEFT FEMUR OF VARANUS SALVATOR. AMER. MUS. NAT. HIST: Pie BaePiGe4tGy | Sine AB! Antero-posterior diameter of bone, 11 mm.; lateral, 10 mm. Antero-posterior diameter of medullary canal, 7 mm.; lateral, 6 mm. The medullary canal is full. Medullary index, 51%. Structure.—The section is composed of concentric lamelle with long and oval lacune and bushy canaliculi surrounding the bone, with the exception of the posterior wall. The lamelle are partially separated into lamine by con- centric rows of long lacunz placed end to end. Numerous short canals, parallel to each other and radiating from the medullary canal, cross the lamelle. Around the canals are clear areas of bone substance crossed in many instances by fine canalicul. In the posterior and inner wall the concentric lamelle are displaced by canals and oval lacune with bushy canaliculi extending from the external surface to the medullary canal. The canals mark the locations of future Ha- versian systems. Fragments of lamelle surround the medullary canal. In the No. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE 5] posterior wall a little cancellous bone appears. ‘The bone shows an early dif- ferentiation by its traces of lamine and Haversian canals. Type I-III, Ia. RIGHT FEMUR OF AMPHIBOLURUS BARBATUS. (AUSTRALIA, ) AMER. MUS. NAT. HIST. 12, Bi, Jae, A Sway, MUAY I Antero-posterior diameter of bone, 3 mm.; lateral, 4 mm. Antero-posterior diameter of medullary canal, 1.5 mm.; lateral, 2.5 mm. The medullary canal is full. Medullary index, 49%. Structure——The section is composed of concentric lamellae with long and oval lacune and long, straight canaliculi surrounding the medullary canal. No differentiation of structure is present. A large, vascular canal appears in the inner wall. The section is uniform. Type I. LEFT FEMUR OF VARANUS ARENARIUS. AMER. MUS. NAT. HIST. Pr. 3, Hie, 48: Syn. Tas, il Antero-posterior diameter of bone, 4.5 mm.; lateral, 3 mm. Antero-posterior diameter of medullary canal, 3 mm.; lateral, 2 mm. The medullary canal is full. Medullary index, 45%. Structure.—The section is composed of concentric lamelle with oval lacune and bushy canaliculi arranged around the medullary canal. Many small canals traverse the walls of the bone radially from the medullary canal outward and from the external surface inward. There is very little differentiation of struc- ture. The section is uniform. Type I. RIGHT FEMUR OF VARANUS NUCHALIS. MONITOR. AMER. MUS. NAT. HIST. Pu. 3, Fie. 49: Syn. Tas. II Antero-posterior diameter of bone, 4.5 mm.; lateral, 5.5 mm. Antero-posterior diameter of medullary canal, 3 mm.; lateral, 2.5 mm. The medullary canal is full. Medullary index, 88%. Structure—With the exception of a small area of lamelle along the medul- lary surface of the posterior wall, the section is composed of concentric lamelle with oval lacune and bushy canaliculi, interrupted by a large number of short. radiating canals around which are clear areas of bone substance. In the pos- 52 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL, 30 terior wall the canals are circular in cross-section, elsewhere they are long. The bone shows a little differentiation of structure. Type I-III, Ia. RIGHT FEMUR OF HELODERMA SUSPECTUM. GILA MONSTER. NO. 583, AMER. MUS. NAT. HIST. Pr. 45 Bie. 50: Syn. Las, Uf 9 Antero-posterior diameter of bone, 3 mm.; lateral, 38 mm. Antero-posterior diameter of medullary canal, 0.7 mm.; lateral, 0.7 mm. The medullary canal is full. Medullary index, 5%. Structure-—The section is composed of concentric lamelle having oval and long lacune with bushy and straight canaliculi indistinctly outlned in lamine. The external lamelle show long lacune and rather straight canaliculi, the re- maining lamellae, oval lacune and bushy canaliculi. Around the medullary canal is an enclosing ring of lamelle with lacune and long, straight cana- liculi. The bone shows traces of differentiation into the three main divisions —external, a central ring, and internal circumferential lamellae. There is no trace of an Haversian system. Type I. RIGHT FEMUR OF SCELOPORUS CLARKI. No. I 61 17, U. Ss. NAT. MUS. 15 2 Ate, Sil, fay, MS ION Antero-posterior diameter of bone, 1.5 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.6 mm. The medullary canal is full. Medullary index, 12%. Structure.—The section is composed of concentric lamelle with oval lacunse and bushy canaliculi surrounding the medullary canal. There is no differentia- tion of structure. The section is uniform. Type I. RIGHT FEMUR OF SCELOPORUS SPINOSUS FLORIDANUS. No. I 73 12, U. S. NAT. MUS. Pr, 4s Ries 52. Sano Dap wt Antero-posterior diameter of bone, 1.5 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.5 mm. The medullary canal is full. Medullary index, 12%. Structure.—The section is composed of concentric lamelle with long lacune and straight canaliculi surrounding the medullary canal. There is no dif- ferentiation of structure. The section is uniform. Type I. No. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE 53 RIGHT FEMUR OF SCELOPORUS OCCIDENTALIS. No. I 74 3, U. S. NAT. MUS. ig, hha ie | rang Me al Antero-posterior diameter of bone, 1.5 mm.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.6 mm.; lateral, 0.5 mm. The medullary canal is full. Medullary index, 24%. Structure.—The section is composed of lamelle with oval and long lacune and straight canaliculi surrounding the medullary canal. There is no dif- ferentiation of structure. Type I. RIGHT FEMUR OF SCELOPORUS MAGISTER. NO. J 71 2, U. S. NAT. MUS. 1h, ak lines Be Swain Abie JUL Antero-posterior diameter of bone, 1.5 mm.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.5 mm. The medullary canal is full. Medullary index, 19%. Structure.—The section is composed of concentric lamelle with oval lacune and bushy canaliculi surrounding the medullary canal. There is no differentia- tion of structure. Type L. RIGHT FEMUR OF CYCLURA CARINATA.- U. S. NAT. MUS. Rie ieee one) SNe ABE I Antero-posterior diameter of bone, 5 mm.; lateral, 5 mim. Antero-posterior diameter of medullary canal, 5 mm.; lateral, 3 mm. The medullary canal is full. Medullary index, 56%. Structure.—The section is composed of concentric lamelle with oval lacune and straight canaliculi surrounding the medullary canal. There is very little differentiation of structure. The section is uniform. Type I. RIGHT FEMUR OF ANOLIS GRISTATELLUS. NO. L 15 13, U. S. NAT. MUS. Pr. 4, Fie. 56. Syn. Tar. Il Antero-posterior diameter of bone, 1 mm.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.6 mm. The medullary canal is full. Medullary index, 338%. Structure-—The section is composed of two rings of lamelle, external and internal. The external has long lacune with straight canaliculi and the internal, oval and round lacune with bushy canaliculi. They are of nearly equal width. There is very little differentiation of structure. The section shows the twofold division. Type I. 54. SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 39 RIGHT FEMUR OF CROTAPHYTUS COLLARIS. NO. J 21 12, U. S. NAT. MUS. 1B, Ze Mae, Big Sway WU IE Antero-posterior diameter of bone, 1.5 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 1 mm.; lateral, 1 mm. The medullary canal is full. Medullary dex, 80%. Structure.—The section is composed of two concentric rings of lamelle surrounding the medullary canal. The external ring has long lacune and straight canaliculi, the internal, oval lacune and bushy canaliculi. The section shows the twofold division. Type L. LEFT FEMUR OF CROTAPHYTUS COLLARIS. NO. J 32 9, U. S. NAT. MUS. 1m, Zo dacs ise Syase dha IU Antero-posterior diameter of bone, 1.5 mm.; lateral, 1.5 mm. Antero-posterior diameter of medullary canal, 0.8 mm.; lateral, 0.8 mm. The medullary canal is full. Medullary index, 40%. Structure-—The section is composed of two concentric rings of lamelle surrounding the medullary canal. The external has long lacune and straight canaliculi, the internal, oval lacune and bushy canaliculi. The section shows the twofold division. Type I. RIGHT FEMUR OF AMEIVA EXUL. (CAYA DE SANTIAGO, PORTO RICO.) NO. L 21 11, U. S. NAT. MUS. ire Ze dhives GL Swan 200g, 10 Antero-posterior diameter of bone, 2 mm.; lateral, 2 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.5 mm. The medullary canal is full. Medullary index. 33%. Structure —The section is composed of concentric lamelle with oval and long lacune and straight canaliculi surrounding the medullary canal. There is very little differentiation of structure. The section is uniform. Type I. RIGHT FEMUR OF EUMECES FASCIATUS. NO. H 31 4, U. S. NAT. MUS. Pi. 4, Fie. 60. Syn. Tas. II Antero-posterior diameter of bone, 1.5 mm.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.6 mm.; lateral, 0.5 mm. The medullary canal is full. Medullary index, 24%. Structure.—The section is composed of lamelle with oval lacune and bushy canaliculi dimly outlined in lamine. There is little differentiation of structure. Type I. o or NO. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE RIGHT FEMUR OF SAUROMALUS. No. J 40 6, U. S. NAT. MUS. Pu. 4, Fie. 61. Syn. Tas. II Antero-posterior diameter of bone, 3 mm.; lateral, 2.5 mm. Antero-posterior diameter of medullary canal, 1.5 mm.; lateral, 1.2 mm. The medullary canal is full. Medullary index, 32%. Structure.—The section is composed of concentric lamelle with oval lacune and straight canaliculi partially divided into concentric rings. A vascular canal is seen in the posterior inner wall. A narrow ring of lamelle with long lacune and straight canaliculi surrounds the medullary canal. Type I. RIGHT FEMUR OF GERRHONOTUS GRANDIS. NO. I 22 4, U. S. NAT. MUS. Pie 4a IGS Gee SIN. SLAB LL Antero-posterior diameter of bone, 1.5 mim.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.4 mm.; lateral, 0.4 mim. The medullary canal is full. Medullary index, 11%. Structure.—The section is composed of concentric lamelle with oval and long lacune and straight and bushy canaliculi. Around the posterior and lateral medullary surface is a crescent of basic bone substance with round lacune and bushy canaliculi. Type I. RIGHT FEMUR OF PYTHON REGIUS. PYTHON. (11$ FEET IN LENGrH— DIED AT WASHINGTON ZOO) Pic, 4 Ie, (ai, Sasi WAS IU Antero-posterior diameter of bone, 1 mm.; lateral, 0.6 mm. Antero-posterior diameter of medullary canal, 0.4 mm.; lateral, 0.8 mm. The medullary canal is full. Medullary index, 24%. Structure.—The femur is rudimentary. The section is composed of lamelle arranged in a peculiar manner. In the anterior wall they are arranged con- centrically around a semicircle with a short radius. In the lateral and pos- terior wall the lamelle take a long curve from the medullary surface of the anterior wall. The lacune are round and the canaliculi are bushy. The anterior wall is best developed. A narrow ring of internal lamelle with long lacune and straight canaliculi surrounds the medullary canal. Type I. 5 56 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 39 LEFT FEMUR OF THE SAME PYTHON REGIUS. U. S. NAT. MUS. Pu. 4, Fic. 64. Syn. Tas. IT Antero-posterior diameter of bone, 1 mm.; lateral, 1 mm. Antero-posterior diameter of medullary canal, 0.5 mm.; lateral, 0.4 mm. The medullary canal is full. Medullary index, 28%. Structure.—The section is composed of two rings of lamellae, viz.: external and internal. The external is thickest and is composed of lamelle with a few round and oval lacune and bushy canaliculi. The canaliculi of the posterior wall are long and straight and, in the anterior wall, infrequent and bushy. A narrow ring of internal lamelle with long lacune and straight canalicul sur- rounds the medullary canal. Type I. LEFT FEMUR OF ALLIGATOR MISSISSIPPIENSIS. ALLIGATOR. CR. MED. COLL. Pu. 4, Fie. 65. Syn. Tas. II Antero-posterior diameter of bone, 17 mm.; lateral, 15 mm. Antero-posterior diameter of medullary canal, 7 mm.; lateral, 6 mm. The medullary canal is full. Medullary index, 20%. Structure.—A thin cross-section of this femur held up to the light presents a ringed appearance like that of a cross-section of the trunk of a tree. The section is composed of three concentric rings of lamine with long, nar- row lacune and straight canaliculi, alternating with four concentric rings of bone substance enclosing crude Haversian canals. The Haversian canals are round, oval, or irregular in shape, are large and small in size, and very numer- ous. They are surrounded by clear areas of bone substance and many fine cana- liculi from concentric adjacent oval lacune pass radially across the areas of bone substance into the canals. The (Ib) stage of the Haversian system is repre- sented. The lamine are fairly well developed. Type I-IL-II], Ia, Ib. FEMUR OF CHELYDRA SERPENTINA. SNAPPING TURTLE. CR. MED. COLL. Pr. 4, Bie. 66. Syn. TAB: IL Antero-posterior diameter of bone, 8 mm.; lateral, 8.5 mm. Antero-posterior diameter of medullary canal, 1 mm.; lateral, 1 mm. The medullary canal is full. Medullary index, 1.5%. Structure-—The wall of the shaft is very thick, proportionately, and the medullary canal is very small. The femur is nearly solid. The section has four concentric rings of lamine alternating with three concentric rings of bone substance in which are many Haversian canals. The canals are much more regular in shape than they were in the alligator and are surrounded by smaller, NO. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE 57 clear areas of bone substance. Oval lacune are arranged concentrically around the boundaries of these areas and bushy canaliculi pass from the lacune to the eanals. The lamine are much better developed than the Haversian systems which have reached the (Ib) differentiation. The posterior wall is composed of bone substance with round lacune and bushy canaliculi arranged as a crude cancellous bone and merges into the cancellous bone of the medullary canal. Type LILIII, Ia, Ib. RIGHT FEMUR OF TRIONYX SPINIFER. SOFT-SHELLED TURTLE. NO. 2325, AMER. MUS. NAT. HIST, Tits 2 IME Ore | Siatig ANNE IO Antero-posterior diameter of bone, 3.5 mm.; lateral, 5.5 mm. Antero-posterior diameter of medullary canal—eancellous bone. The medullary canal is full. Medullary index, 0. Structure.—The section is surrounded by a ring of lamellae with oval lacune and bushy canaliculi. It is interrupted by Haversian canals of the (la) dif- ferentiation. In the posterior wall here and there an Haversian system appears in the (Ib) stage of advancement. Cancellous bone occupies the central canal of the bone and is derived, by extension, from the enclosing lamellar ring. The walls of the cancellous meshes are composed of lamelle with oval and long lacune and straight canaliculi. There is no individual medullary canal. Type I-III, Ia, Ib. RIGHT FEMUR OF CINOSTERNUM PENNSYLVANICUM. MUD TURTLE. AMER. MUS. NAT. HIST. . Wim, GE Me asl, Siang Mi IU Antero-posterior diameter of bone, 3 mm.; lateral, 3.5 mm. Antero-posterior diameter of medullary canal—cancellous bone. The medullary canal ‘is full. Medullary index, 0. Structure.—The section is surrounded by a narrow ring of lamelle with oval lacune and bushy canaliculi, from which is derived the cancellous bone which fills the medullary canal. A few Haversian systems of the (Ib) dif- ferentiation are found in the posterior wall. Type I-III, Ib. RIGHT FEMUR OF CHELOPUS GUTTATUS. SPOTTED TURTLE. AMER. MUS, NAT, HIST. Pu. 4, Fic. 69. Syn. Tas. II Antero-posterior diameter of bone, 2.5 mm.; lateral, 4 mm. Antero-posterior diameter of medullary canal—cancellous bone. The medullary canal is full. Medullary index, 0. 58 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35 Structure.—The section is surrounded by a narrow ring of lamelle with oval lacune and bushy canaliculi, interrupted by a few Haversian canals of the (Ia) differentiation. From the under surface of this ring is derived a ‘ancellous bone which entirely fills the medullary canal. The walls of the meshes are composed of lamellae with oval lacune and bushy canaliculi. In the posterior wall are a few Haversian systems of the (Ib) stage of development. Type I-III, Ia, Ib. LEFT FEMUR OF CHRYSEMYS PICTA. PAINTED TURTLE. AMER. MUS. NAT. HIST. Pu. 4, Fie. 70. Syn. Tas. II Antero-posterior diameter of bone, 2.5 mm.; lateral, 4 mm. cancellous bone. Antero-posterior diameter of medullary canal The medullary canal is full. Medullary index, 0. Structure.—The section is surrounded by a ring of lamella with oval lacune and bushy canaliculi, interrupted by a few Haversian canals of the (la) differentiation. Haversian systems of the (Ib) stage of development are found in the posterior wall. From the lamellar ring is derived the cancellous bone which occupies the whole medullary canal. Type I-III, Ia, Ib. RIGHT FEMUR OF AROMOCHELYS ODORATUS. MUSK TURTLE. AMER. MUS, NAT. HIST. Pu. 4, Bie) wl. sya. ABs Uf Antero-posterior diameter of bone, 1.8 mm.; lateral, 1.4 mm. Antero-posterior diameter of medullary canal, 0.3 mm.; lateral, 0.38 mm. The medullary canal is full. Medullary index, 4%. Slructure—The section is surrounded by a band of lamellae of various widths. The lacune are long and their canaliculi are long and straight. Under- neath this is a central ring of lamelle with oval lacune and bushy canaliculi. In this ring are several Haversian systems of the (Ib) differentiation forming a circular row around the medullary canal. Around the medullary canal is a ring of internal circumferential lamellae with long Jacune and straight canaliculi. The bone shows the outlines of the three divisions—external and internal circumferential lamella and the central ring of lamelle with Haversian systems. Type I-III, Ib. RIGHT FEMUR OF PSEUDEMYS FLORIDANA. NO. 28417, U. Ss. NAT. MUS. Pu. 4, Fie. 72. Syn. Tas. I Antero-posterior diameter of bone, 4 mm.; lateral, 8 mm. Antero-posterior diameter of medullary canal—cancellous bone. Medullary index, 0, No. 3 COMPARATIVE HISTOLOGY OF FEMUR—FOOTE 09 Structure.—The section is composed of lamelle with oval lacune and bushy canaliculi, incompletely separated into lamine which surround the medullary canal. In the posterior wall the crude lamine are frequently interrupted by Haversian canals of the (Ib) differentiation. Im the internal laminew of the anterior wall are several crude undeveloped and a few fairly well developed Haversian systems. The medullary canal is occupied by cancellous bone. Type I-IT-ITI, Ib. RIGHT FEMUR OF TESTUDO (GOPHERUS) POLYPHEMUS. NO. 75955, U. S. NAT. MUS. Pins iG. en aS eNe MLAB TT Antero-posterior diameter of bone, 8.5 mm.; lateral, 6 mm. Antero-posterior diameter of medullary canal—cancellous bone. The medullary canal is full. Medullary index, 0. Structure—The section is composed of incompletely formed, concentric lamin which constitute the anterior, outer, and posterior wall. The lamine are crossed by short, radiating canals and interrupted by Haversian systems of the (Ia) differentiation. Beneath the lamine of the anterior and outer wall are Haversian systems of the (Ib) stage of development and the posterior ridge is made up almost entirely of Haversian systems of the same develop- ment. The inner wall is composed of lamelle. Type I-III, Ia, Ib. VIII. BIRDS Forty femora were examined. GeNERAL CHARACTER OF THE I"EmMuR The femora of birds vary considerably in shape. A few are triangular, many are elliptical, and some are circular. The majority of them are elliptical and their antero-posterior diameters are longest. The medullary contents pre- sent a variable character. In some femora the medullary canals are full of marrow; in some, of cancellous bone, the meshes of which are filled with mar- row; while in others, the canals are empty or occupied by trabecule only. About half of the femora examined have no contents. The medullary surfaces also vary somewhat in character. In those canals filled with marrow and blood vessels the surface is uneven, while in those which have no contents the surface is smooth. In these bones the walls are thin, the canals large, and the trabecule are numerous. The medullary index varies from 0 to 327%, with an average of 159%. The bone structures show considerable variation. The three single types and many combinations of types, in an incomplete or complete differentiation, 60 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE vou. 35 are found. None of these, with the possible exception of the first, have reached their full development. The second is found in a very incomplete and an advanced stage and forms the structure of a large number of femora; while the third has assumed a more complex form than that found in reptiles. The Haversian systems are comparatively large, the Haversian canals are small and around them are concentrically arranged round or oval lacune with intri- cate networks of canaliculi. This is the (Ic) stage of Haversian differentiation and is characteristic of birds. DetraILeD HXAMINATION RIGHT FEMUR OF CYANOCITTA STELLERI AZTECA. AZTEC JAY. NO. 2874, AMER. MUS. NAT. HIST. Pre 5, Hic. 745 SVN. UAB. ll Antero-posterior diameter of bone, 3 mm.; lateral, 2.5 mm. Antero-posterior diameter of medullary canal, 2 mm.; lateral, 1.5 mm. The medullary canal is full. Medullary index, 68%. Structure.—The section is composed of bone substance with oval lacune and bushy canaliculi surrounding the medullary canal. The bone substance is partially separated into lamine by short canals. The bone shows but little differentiation of structure. Type LILI. LEFT FEMUR OF MERGUS SERRATOR. RED-BREASTED MERGANSER. NO. BulIl((, AMER. MUS, NAT. HIST. Pu. 5, Fie. 75. Syn. Tas. IIT Antero-posterior diameter of bone, 6.5 mm.; lateral, 4.5 mm. Antero-posterior diameter of medullary canal, 4.5 mm.; lateral, 2.5 mm. The medullary canal is full. Medullary index, 68%. Structure—The section is composed of bone substance with oval lacune and bushy canaliculi, separated into incomplete lamine by short, concentric, branching canals. A few vascular canals running longitudinally appear in the posterior inner and anterior wall. In the bone substance are a few Haversian systems of the (Ic) stage of development, and a single better differentiated system occurs in the tip of the posterior ridge. 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Ge| Ilh| XxX ” Callens 9T TZ} 02) 92 92 <5 ia Vee | nema | eee [referee |e NES COAT OST: ” » ‘oTeulay Gé| OlF| X ” males 9F |G'ST|G LT] TE v | X os X|X/T/ OND OOF » ” n | G&| 60%} X ” mien 61 GS ee | ees 8@ XS || 8 X|X|xX| » TST » » » G&| 80h) X ” ON PAS LE ST 9T 6% T& x é SSelbes Ila » 691 ” » ” Ge| L0b) xX ” CON x 06 TL IL| 9° &@ Té SNe 4 x| a ” 9ST ” » ” GE| 907) X ” elles G& | STE ST GG 8% es forename || Xx | 1) ” 69T » ” ” G&| SOP) X ” OP liens 86 ST ST | 9°82 86 x - x | UW) O° N ‘CW 09T ” ” » G&| POF) ~° ” x 06 OT FL | SLs 06 x xX|xX|xX/ 1 ” 66 ” ” ” GE SOF a? ” x GP GT 9T LE G 86 x : x x x } ” 16 ” ” ” Té| 20) X ” x && tT LT 64 Ts x 9 1X} on | ONAL OD 0&6 ” ” ” T&| TOF) X ” x 0& FL |S°&T && iG x “|X| | ON ‘GW 871 » ” ” TE§| 006) X ” Xx &@ AB) IE 26 Oi eel WO] eas || nO UNTO 7X4 » ” ” Tg| 668) xX ” x 1G BE tL 92 |9°Le OS Ne SENS] SNe tll ” SST ” ” ” TE | 868} xX ” emai 96 Cm Kayes 1s CYA eon h a eel ete [cage Soa] [onl ed ” TOT ” » ” TE | L686) X ” x FP ey lies) 63 0€ Xe | I| ec |ma | a > oI t | ow ut | al z 3 Ble) 72 # | 55 its E ale let och eure yy calles & = g ce @ | Bs = gs | 28 oF s SstIO ae fs BB eae es cee spel ee Sad ee | oe ee st = =P = Sf = eo se = Baa leseae lee Sule Ble | 58 Z ae z 3 5 Z aoua1aJoy [euro AiR [NpaW F = 5 a [euro Are [paw auog ja Spiele wnsEa -1quioo mao sadAy, X WZIAVL OLLGONAS VoL. 35 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE 230 ‘UMBIP JOU ‘paqiiosed » semen on esp GSP x x ” x gg| Ish) X ”» SEN 1 lal) Vaz alkenes x ” Dales 8& SHE Jt pill a x ” SN es 1Z 6 as si | dene x » alas 96 | LT eal et x ” => ess GE ARI AI G&| Shh] Xx » Pes DEM Sach) esi Pall x ” TO eS 0€& |S'&L| OL ills x » x 61 OT) &. 54 ees + ” LPS 62 tL eet SA eon x ” ales 8& vr) 9T ei|* 24 ” x cg Shee GE) 68h] xX ” ilps 82 WEN) Mat SAR x ” ales OOT|S°2T) 92 Ge) Ler) Xx » SE Wes 0g FT| 60 GE) 98h | xX ” Nes 0§ 9T| ST ade || Rice x ” melleN 63 Gh SE Saullo sa ” x 9% Go| Lz Sol ” allies Gé GT} LT oy 2% ” be x 63 &T $L elle me ” Tex *S 0g; L- we . x x x 1X KXKXXKXKX wD co sno|joouvo De 19 a = oO a > SPS 61h STP | LIP OTF STP FIP x ” ” x AY xX iXKKXKXXXXX S& | &IP ” re Tg Ot |OL /|8. ee ig: ” ‘ 19 Li OF €&| GIh) X | Snoy[eouRvo 06 9T| #E) Le | | ~| Ln a | = Is >| oo Seal z Ble| ye | 2 | se] & ® 4 ce 2. 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(pozejndure) ,, Ok ‘DW OD 89Z Daan (OSECSE) nas, 1» & ” § ” aha phe ete ” ” ” | ‘AO 'N ‘a ‘NZ "ON 82" (49TATIOD) ou ‘Ore | ” ” ” 480} eB] ” ” ” ” ” ” 19} ” ” ” ” | Ayres JO 93v4s A[Iva ,, teeeeeeeess sepTiauiel SUIMOYS w94s4s UeISIOABHT e1surg ‘NN ‘CW GFL ” ciples cena AC OP 93v) ” ” ‘OO #6 Dit tteeteeeeereees “OM |, WN ‘N'S ‘QM 0¢pLzz » coc uerpegsny rs T6 ttt tees eessereee ee ** (epyomns * va 8 age) oun ‘ele ” 99T rhe es * POL nA . re a aqTy a ‘gpeulayy } a 7% FeO CSSONGGO0 (‘qo00s | eAoqe MO[Eq “UM ge) ,, | ‘9081 ‘1 'N ‘CW GST ‘ON’ **** (ANtey OTQNOp) 931A ‘OTe | OFPIM uN | ]BULUTe yy ss8tID IX WIAVL OILAONAS INDEX PAGE Aberrant method of Haversian development.............- eee cece eee te eet ee renee ete e eens 16 Adult femora—types and type combinations.........-.---..-0+es sere eee ee eee yap ae Be T2182 ot UNGHAIRGES) Tin Tees) tay iesoon soo banboeaepoboomy poo os OF Ubu buo cE Go0U 0D dbcoboeDCosoesGDo bon GunC 13, 16 Age, influence of, on bone structure.............. 2 eee e eee eee eee eet eee ees 25 MONDE Coossn secon sod sbaoucdnooss Seen cose CO ORD On0 OD opamp ouoUD cocoon UdKOgonCoOUC 35 Amphibian type Of bone. ... . 2... 5.5.2 e cece ce ee ene ee cen tenets nec tisis ee eeeeccine 35 JMG IBPARSHEN. 35 65nnob0 pao b bo Hbeapono doo coeHodoUbpe Hoon. apc oUnDr a Commmon ccc UO oOUI UIC 177 Atrenaren medtullarye AMCIGES! cic). oar sieve o efole imine ©) airmieinis tielsiaa)minjalevels mime aieinl=inialsia)afels)s le(eveieyals 6 Basic bone units......... Ti poate Ren Sen ee ee en an Gaon aG mets r ao ashi Ao telin Ot ic 112 Basis of the general Summary............ 6... eee e cece eee tenet ete rete eee 3 EDIE) oo¢enss dias as bobenn pase R aU UO ee ADUDOD CU Cone como nOoCoCo Poo ODnT Deron Sano Cima conor nc 20, 75 Bin: ooacscdst ce Uda doctor ooeenocnenomabaccoco don ued cu atpontor DOU DU aetE go SoBe UM OD aos. oCK 19, 60 Biedk RIGB. coda cocoon bocénbeasuuadn pote lo DOdee cor cad Hon dc ocuUGoTHocUonO Daa ONoo aod hs PD. Bhi} T1583 Bye GK socgcudoosmbacdese dod puucds aa ornactd Co peOObon Abdoou TD OU ComunnoDnOsOnop a oncodo 10 (G) stage of Haversian differentiation..........--0.+ 12-12 s esse eee eee e ets tetas 16 Gaciherih so oeaneduaandeoos cpa sen sano cd00 euereeme Op ODO DO CoD dOnOnuO uO Sood o Gt COIR oIsOt 10 G@ancellous bone and trabecule.....-.... 2-2-1222 eee eee enter ets ree tte e sce ee 9 Changes which occur in Ritts Po cane Ub Some cane AAP RO IABOS Bc ou gad OCnoruotocuResBonnopHoDede-y 3D @haractenioh medullary, Surface. «a2 < 566 cece ae eee faerie wir t wm welt imines elem ells ose 8 Garpinatien TEM, “jogaed foeeoksiseeeeee bo cobop oes saben os Nr GonoDoo Uo ORC Ono ODUMOR OOO CPOE aD 18 THIN, TAs Tn, 1, © aaeocoooceosseaoenano cos Contours cacDoUSoPocerSUncCOSEDmOKDS 18 MEO TEs Tis WEG Gooosdcndéecobsone go sono Ssecun Doo Ie o0Re on oD Reno a OOSOnoUC COE GL 18 IRIN, TR, Ty WO C! Sosccocooenn so scoocadoe commr Coupe cod pH pus aUe OOF Onno apo ST 18 Complete differentiation of the Haversian system .........-.22-cee- eee secre tence c tse 16 Iihbe) ~Gonuenoens shane oodbalodbeorashn ntpouDsOddonooRcnaS 14 CimnGkisieRe sss dsc Jegnecee Bano COU ae apne cana coe Ome Samo OF OOo mom mOr LOOT aD ODO ORC Od: Par, Montentenotathemmedmilary: Canal cer. istecs aias cle eae ier sors levels olde ohne rio eserminin ey sie) =) -mirasieseeler= onaieinvelainie 8 DAiniion Ge Tee wOccia) Gh Alieuth gpg onppon ee oodoboencd op ued oanpop oO UOC Uma aon S 20S OG ary ala! AGNSIEN. cSanceeagsoeeun’d dud dpouepos Doo OUe pou carn pobgmnUoy SoOUb OPA BIC ao rag nic 10 IG, Ss anocasadeb noc nuder a woe oder er cn oOo no obdaur cnousOn oo reRa Imo Od COS Te 11 IDAGRS 26, cccn ene dos aeneOneeeaeheb Conc Une BROd ee no8 = acOO I reo Opi ULEIT OD OC Sr Gib E ae. 10 Dwercitay OF PONS .sapscssesodeocsoon one acon A ande poo goes) GoGraDUnD pon CHAC CSSD CH OOS S 5 Mescriptionmot firsts type WOW eye -ieta (ae neler mreelmaslerein ciate elo liens hele ame ag 12 Eicemul ian) Wane) ppasocuadosom ado00 se counoOpo DOO OODOOT: A etre tayaheys commuetst=ea= 3 sinimal, (aioe) ING) osedonsamoasue cde aesooo Dono ScDOCNondtaundOOm eh DOUNGESoOROr 14 Metailedwexaminationwor amphibians.) c-.-1-%+ >.<) - jo. ciemie oral = ane iain ciel che ons aoe hha SS a 35 TINH ES ago One U DRONA on wase aon CO UMOREApo Gao yR MOD Adahdd ICIS Aca 48 Nils) sanoosmooeSoospasooe rn SRE RIA ods Gai cio Ae OSM IF ORM OCOCaROC 60 TD Sdopouncdaescuscg co ee.unn ccobritD oc BasnOn ond OisiySOc Da On nis iC 76 Ging TMT: Gooenoatans DOU Ret OU cOD ODOM OUEOn abc mar ratel iottices 90 MEIN) SaSin Gunsodeaw CF ONCE De CELE Ome ace AUD Dior wigs Ser OOO OCHRE DOH, 149 Mevelopmenton tievlinea: aSperae jie) -lacimi= sss Sores cies my a Sinclar ite 30, 31 lamellar and laminar horseshoe-shaped ringS..-------+++-+-+++rerrsrertete 30 Developmental stages as seen in Pueblo ImGkGnl a) So peneaeo eH aoe SoU CeO UC ODOC ood GoDG 31 Miterentiation of Elaversian SYStCMS ~~... -----eccsc ere ere Ts a 14 iGHAGIER aouccaasasss OO be OEE nS CHCOn eer anon Dao cramp CO GCr ROG AOR OC DOL 12 GHD Di dooa aco b CO n GOCE BD abe OOO OO AEE IO TRIO OR or get cease raiat ag ie 14 TOYNEy HTC 9 Oak oe eho RE DOSE Bee a2 tin COG R mane peas OOD ReY cc ari SC CIOO DHE 10 iA) GSES) god Sucl/eon SEB sees coon Boe adn cOpP an comMonbm the mEnio Jer DOS SOG 12 Dieeane, imines wr say acess snbaconcsgenbosese 2 te anoOm ae BO Dr COE sgk aS sg im ly 27, 212 Distribution of the three bone types as they exist in the various femora....----++++++s+sseee: 17 Egyptian femora of XII Dynasty ......--------+-s+esestrs rete Re Fei rotors srotetehetagsiieer= 23, LTT 16 232 INDEX VOL. 30 PAGE Elliptical—Most prominent shape of the amphibian femur ...........-..:seeseeseeeeeereees 4 MA HHVEHOY MAhIbe Somos KomddsosodGUdoMDODUnoOOodaoOdS 4 bird! Heme! a sieve cs ca.2 avs = fovetets overeat ayer ole ate wetetotaely a) aieiatar a 4 DatiwhemMiUT ye) jerearere acta eee reke eepdes elec ec creesietereperete ite 4 mammalian: Tempy 6.05. ko istereus: «arate detees = )een ere inieysatav halal 4 Femur PELOWAMAN Sa, wr parser cieay nerarele ate eetoreh teeiel sister = 4 Bividences: of advance im) amphibians! sor epee rete cre let ncte etc haee) eet te ate tata oan alee telat aoe tet os eat aY=I 18, 29 Teptilesy fsicpscys srateee cuanavs ate se sustapevatececere t= else te cerekoleroteliotetete eter let ectstenee rar = tel aS PAs) Lt) 0 oi |: enc micCricn cic coc rata eho at titrate = 19, 29 Bets occa SSS sya aes axceadilay as ene be ee eee ecisbanr sabe esievege ane lara alt care alee onaiebs apart 205 29 other mammals) = iy. sa facie ness eee eronein tae Le ieee tel ee oasteeite Cmeao WMD ods Sisicretec. cafe, deelSck. 5, sherassrer Grate te chaka abansts levees Rerishete te etehorsrecetatte este eat timate te geramaterEe iano) type ad VaNnCementy 2.6. wietare cvsver ermine a shake clr ehe eave et Moioes ele istsie cael etched ieet renee tensions PAE! HXternal Shapes: OL LOMUOrey eer. creievelciel elects ratstenaterasctclctaite) orersretel elas porch ataterede = patretta tastes fatiete tethers ieteee tate Tai 4 Factors influencing: bonestype of StUrwctire, sscc celeste severe ais tee ioe ete eae hetero a weet 24 Pemora. Oxamined) .<2..55 acc co csegs rere Ceo eens edits ase re SUISSE Te ears ee Tele att oite eae One RRC OTe cee tine ete 1,22 OF -AMpPHiDIAMS: -os.0.5 55 | eis esata Sete ete Gea hvew ye eee aaah ase er shsanhe anecepmesyane seta el aieral enstetetmtte eames aiers 34 r<(2) (G18 (0) 0 et 0) Cae al ade a aoe ee Ie nts Sis AIA OS. OOD Ob OC o.oo Citar cae 2 Hemur’ of: SACwIS: STV US! oe terol ce eis oe Sesto eutee wcste ene elke eer dette eae Rete et ee see an se eee eet es ee 41 Ajaia “ajaja. (roseate Spoonbill neva centre dorcier er Tonete ota cate teva ee taad atest 61 Atees smachlis; (Gel) sis. 5) ros ls SiSie oe ree ereasrorckcte a che nett ee ee ee eee eee nce 105 Alligator MiSSISSIPDIEN SIS? ac) domauoe shoo tupao DU nC 138 (OHMmhvOre TmG@rane (Glee) sgncauasnesnsoooocbouvosednooGocongdUonUaooS oD oOouCUdTS 105 Gastonicanadensish (DEAMET)) sierys eledeccieye =) ctelriele era ailsielmieiats (olsviehel«leiiokete)-¥-r=)a1auc)atanaleleteleteelna 125 Gathartes aura (turkey-buzzard)) ..-.. 522-266... cee eee ee ee ee eee see 75 Chany Qniksal (fnicemnots)) sesegnoee aon ce oo oo ododoododnG on ooobo moon oeondonondor 92 Centrocercus urophasianus (sage hen) ...........-- eee e eee eee eee teen eee eee 66 Cephalophus (African antelope) ......--..---.s see e eee teeter ees 136 Chameleo vulgaris (chameleon) .........6... ees eect tee tee t eet eee teens ; 49 Charadrius pluvialis (golden plover)............... sees eee eee eee eee eee eee 62 Chauna cristata (crested screamer) .......-. 2. cece eee ere eee tee e ett eee 70 G@heiromeles torquatus) -.<.5 <5 52s 2c ce ee ce ee pee we ems ies lee ee wien meen tele pe 87 Chelopus guttatus (spotted turtle) ........... 2... se ee eee eee ete eee ees 57 Chelydra serpentina (snapping turtle) .........-..--- ++ esse eee eee tee eee eects 56 Ghicama Indian, No. 2, U. S. Nat. Mus. ......... 0.0 cee eee cece rete ete e ne ees 170 3h Wh Gb ING NIKE Aondooocosee cupomaoua: aede conanddbodocgar 171 1, 10S SO INET WEE Sobbourenocpaeoodc baduosoupcconronsoooOS 171 7s Oph INE) UNE one Oo booed eon ecosoR ooopddadonne Hoses 171 Th We REINER WMGb» sacnacsosenpoosoaod07 puonens sandagDba050 172 OU eb aNehn WONISh Mesa codbacousadascumodcrerosA= nab eouCoUGD 172 Gilles sgoeccogossccs JoduD ooo odor AH JURE roo UScOoo DD encronoRe gcc anon one he 85 Cholepus didactylus (Ciimoaxeciel oid) Gaaahs secouooD Oe Scnood ans eboovece NCD Gob aGncr 109 Ghorophilus feriarum ... 2. . 2-2-2222. ee ee ee teen 41 Chrysemys picta (painted turtle) ......-...- +... esses etre eee eee ees 58 Cinosternum pennsylvanicum (mud NAN its Mobo nt aobhGdo she SOOO UeDeTenoChgs tok ao 57 Colobus abyssinicus caudatus ........--.+- 2. eee eee eee ete eee tee eee eee ees 94 Connochetes taurinus albojubatus (gnu) ......---- ++ sete eee e eee eect eee eee ee eee 107 Corvus americanus (American crow) ...---------ssee eee teers tees tees 72 Grotaphytus collaris (left) <2... <2 5222202 - 2-2-2 nn asain 54 (Gaia: Rens ooee sao cmeaMbee shou Ads ceoocdaedennadodupupdia? 54 Cryptobranchus allegheniensis (Una SAID) Goeocs sodas sacancn gd dessa nen or dos oes 3 Cryptoprocta ferox (cat-like civet)........--. 1. seers sete epee ete eee e eee 118 Cyanocitta cristata (blue jay) ....- 2.2.2.2. este ee te ne eee eee 62 stelleri azteca (aztec jay) ..... 0-22. s eee e eect cee rte teense tence 60 Cucina, Came, sca kcesdo cmon anens sposeeooer ee sro Uacd Ubu setor iGnePcUrchanaoBe 53 ChaGuAlng (elite) 4. coecgease°osocnsesurOn > s0o0 G6 bb oOoR be pEduacoo cy Scarocr 101 maimon (mandrill) .....:... cece eee tet eet eee ce eee ee ene teens 101 Cynomys ludovicianus (prairie dog) ...-.---++-+2++ +e sere reer eerste sete teen ness 96 CHMONEAS scoucsqconagoooneasoogseDar cs Wnondans 9OOGoscdoAmbIDORRegboo bo OR pOOE 86 DEENA BONN! (EVSONN)) Geoceds odo saeneree Os SUSoDG ORR adn odu one cocoqscse Ib: 119 DEGREE) senGebINS! Goosopssnebess4a> 8 sa> cod bos o DUO od aa aon colconeeGaboduIc 81 Dendragapus ODSCUrUS, (SLOUSE)| = se i aa ear wa nian aia oaaaet 67 TDRSS) NCO) Guasasopocdasnone 500.05 c0 gE SmKCRROU ada Geo soc Ag Osc ao 00D 40 DERG TONED oonebosoonsaocsdnon colo dunn ndetOnDoOEGbo oc cr oady oO rucadanor 77 Midelphis vineimiama (OPOSSUM) eae leslie gees ts ele 147 (DHS gortonsenecoe doneuacodg ede shoubas Gt.o ope Sen OcES On BopOnms Vora cers Po SiO 139 TNO EAN REGIE Cline ree acts ene 29 fa se eicaeyerm eeieleralaeinn= oie ere «teria riue ee Ale ae 140 Dromeus nove hollandie (Gui) apoesscooceocgcnud oon oUpUODo OccraCoosono I BoGooor 69 Echidna (egg-laying AULEUTA TAY ULI) gsc teven wins ucts nba ep ats ae olede eke doVeke=le|lescwie lm = /tekerievela|esniiatela eliela\= 91 Meypian) temore Of RIL DYMAStY: ovr. eels dolar eis cine eels a cioinin oh manasa se aa 23 Egyptian child of one year (XII Dynasty), No. 256479 (de), U. S. Nat. Mus. .... 177 256479 (d), U.S. Nat. Mus. .... 178 youth (XII Dynasty), Nos 25Os Oma Lie sos ING MUS! ore a iaro. peepee olnfalsiele ote 179 adult (XII Dynasty), No. 258675 (a), U. Sh INFRRUWINER cpa de Sh ooesepo ass 179 OXMCH (GDS, 10 Sy NETS ibs | ios aes eeacsdeonas 180 ARH (EN) 1h SE INENS MEH Gagbedoscdobodese 180 234 INDEX VOL. 30 PAGE Femur of Egyptian adult (XII Dynasty), No. 258675 (e), U. S. Nat. Mus. ................ 181 ALCS ShaNEhin IMIS pun éoatds sa 2aopos 182 PANT CNC Bas NEN WES Tonmanaanecnndaed 182 Wlephas african ws) Sercktetestelers/-Yatctal=/ Sr API rae oe On Ree tne eek eee ete 103 sheep, D1 weelis {Sas gicvevcte oteraroyotevale hatin ate le Sacre eee Se Coke epee ascot pete tale sheet che reece nen 102 Rox terrier (not, apUre: DIOOG)) ek cec sesh cave se etenie asic ete te ter ae eels tee 140 Galeopithecus) (fiyine Lemur) same sare taieete article ede ieee rie eter ieee eee oe rae 99 Gallus; (domestic chicken) EER San eo ECE oo eR RN eS Beta ema S 80 Wikio pinMlie “7 gopceoos sopooendsnoeeoanoe IS Et ACIS Na MODE Oto ate ee GE Ie. aR 85 WIGAN NE) Wn deo Become REot.ob cata Oko aste sy AO rO ORDO C Gon conta CHO Gna ane cnteme 76 NAAYT Comme SUH ts i ance tos ot = iagcueacvet wiocatcvere lta cicichess AeTe e/apama SG eters SuAe mite felts e. sherm eee set ice 128 CARATS Ae are ARTE ODES CES cng ROLOR IE ore ERNE ROI OREO Orie Sen aS ere 128 POT Ts Se EEO a ORL aE et ea Saat Ce ry eer erat 5 Ecos OIE hs te ee er 129 Oar ea Oe ON MOD Tote ee weec ae A Sten chy ey cae bet PAS Tanenstar Pave omer es Rte ee 129 AVINT SME AC ELL Same mea ter sty eee a cent be eels Gee Ste vacsteie acslche \adlolaMewebeveier et evenGve Wie cars lose pol caret Pannen eto 95 SY VA GLCUS Was top eacte clic ee tetes oe mucvolmcetoieeiacacter Peto sey isle Pane wee sebepaevel eaves tempers nirste Seach ee 98 IMivoralemmoschatan (GeSman Mars cat cies ae a aiecenia serie citi Slat t caterers alae are are 96 WRG TOME” ob otes bod acendbsosde codon sects ogeuensd seonndaoncaoodbadesceageac 82 INGEIEATE) ROME IES (Cie hal)) Gham. noe oon oom onus on Fodms doaoas Done coDcomo nT 45 NGERO, INO, DAES TUS INGE, IMIS, Giggn dodno co onoguuo neu denen vonbdpeedassonerconsc 155 Neeroand white mixed—at least half White... 5. .% 159 Negro, No. 1, Med. Dept. Tulane Univ. .........----+1--- 22 sss cess eee neces eeeee 159 UGhiite Nf, PAM WWE ty INE WE ho6 aaonainecdgeasco0 so Qdcg dem soccecos 160 (right), No. 248674, U. S. Nat. Musso 2... ote ese bee wie os it eal ee 160 (left), “No: 248674), WU. Si Nate MIS ee acne steered ce tara mha tot aoedol=)slelenni=tel 161 IN (ORC YAR ERS Seontioao Cora conrocamcdcets Tod ono m om cdmaricee Sor ouch 164 No. 83, Med: Dept. Tulane Univ. 2.2200 o se eee ee re ee eel 164 (Ay IMME ADloley UBUD Whshh, Soeoeaconsc noose soscpreusooonadooCIss ace 165 63), Med’, Depts Mulame UWintiveg Wernceret-taetersrmtote tere stettor le reatgee itera oe Ro nceetols 165 5, Med) Dept» Dulane: Waniivg ete. cyetretie cvs ecte eden s stot eased alan aias atm a Te olametele tots 165 8, Meds Dept. “Wilarie: Wry ireree t= ara ecto eyed alates een era Lele tedon 166 7, Med. Dept. Dulane, Wmv creer eters here vepatai= ete nor aioe Rer od tolchetaa 166 IN (vob (ANG no Oe ame oO UCno ite COG oDeronADed sn sopro os odose OoOoddSC 87 Numida :meleagris) (euimea LO wily)! sieve c= cyenetese sats tate ayes ste teed cee lok ie otis = feta toe aod etal stele aanie date Polen 61 INAGENIIS CRACKIN wEodeoodromocuasonoop dD. opMocduccosubouneoc Sue oS EOD SSogNOp UL 79 Ny.cteris: DOrealis) “A esice scree cello isle cies) erereteaden chat erated fate fetal etevaret- Reet eeyeeWe ot nate ees etek za 83 P00: (2: ee eee na Bete AoA ORIADIIN OD nino Cla ama mors romom page os Se 3 Nyctherodius violaceus (night heron) ..............-+..s sees ee eee eee eee eee 64 NAKAHKGEME) This! “ooopesonsaconopoonodoues ssuO sD OdDoUDOOSOsUn US oascKSCsenoS 82 (OOP eSB wooo ouaonse dondonoDoGedo0 cadoonoUDdy po abUME.DOddURcooEdouoDIobogdSboCOs 71 Ornithorhynchus (duck bill, egg-laying mammal) ...........--..-.--.+++-+-++--+- 91 (Ghat ofortoalorceche) yehyohia( eshte (0) Soangannohood oo Oooo Do dcndadodgudusavHoNooDAsONS 107 Oviss (domestics Sheep) erate eee ie cisteterieeler ier stare rater a EAA Ba CoG Cp CUdeo OG 127 montana, (mountain SMe) ieee ce syercsetv ete steers elec ate esol of eee ele ete ke et elie Pe rel 135 Pandion carolinensis (American OSPprey) ........5200eeese cece ene nets nee 70 Pavo: (eristatus) ((pearOwl) Ny sere te crete tepeto tal sveneta n= ete dstota alate leat tt ribet ote aoo8Dia0 64 Pedetes: Gjumping Hare). Ose ctee cyeseeteetl- erate, sv avesarereretsialeioverseses os ieee esate ear tah bales liet= Estate 124 Pelecanus erythrorhynchus (white pelican) .............:..--.--+-+-+-+ss-ss-eeee 63 Peruvian) Indian, adult, No. 266469/(b)5 Ws S: Nata Mish ry eerie renters aieiet=neren 170 CAE Me Ue Sh NEE MES Sap soetnaodcsc scar obosds 170 140 5Ue SaeNat MUSH See ees rel roctecerne tent reins 173 ee UR Sy NE ke UOC! en Awe ooo nomecnorcgnouccdcgc 173 Sore Se Nets IMs reir tr ec) veers che ieee eee rate 174 child, No. ihe Oto saeN ts RAN er Oman momcccoanooraonuaacdoe 174 , 19. SS Nats Mis? Wi. Aye fetes erate tena bate rose oe 174 adult, No. LOC GS SS Nate MAss® Geronacc eee Sate ainreieore Anica tere 175 Des Se Nats MUSit sei shgecrsi serene ic rctoleaeteriecetetentate 175 AGO RSE aN Gh aban ecasocdase soem dco. no.c5G 176 Pa 0 em Pees Fel) Cin GI GRIBIeR Scie Boot IA oc tlc Ss 176 Petaliat 28 can steer vais w evesbibioZs uakayavels ats btaue. ahenetele Ter aye Gusvepetanal stoesats leusl cess aes Cae Ve pee Venere ensts 85 Phacocherus atricani ses Gwar, OL) acta teat btersy natal tele ttel hetelat ol oka tee ete eet ele Rete 33 Phascolomys: ursinus: (wombat) yk cree ccces tenth ore ete enne ate ocean ne otra cers ieee one etera teat 97 Phasianus' torquatus: (Chinese spheasamty)i wectercrersietsretspevatatae tetas tale teietaacnal onD coo UOn Demon ono TK . 44 Tae ooocdesesue rose oo One phon puoMmeONDOODoUS SonOa DUOC ODDO Orc cc 44 holbrookii (spade-foot toad) ......--.--+seeeeee reeset ect t etre teres 44 Sadicnmare Gems 5 coscdscaccamdoocacons sponDD Boe IoCODh Gp ODED om Roo One mUCD orc 52 MAURIE osooeusedepoopeacescdsasS ooadopbecuan Da sooo GED dOU GSO DUOO DOC 53 GRMGIAIMEINE coaoosconsnds sD cono0oonDUGon du OOOO Bo mOOnOnDC DS coUboOr 53 SPINOSUS fOTIGANUS 5 = )afre cere «o/c ele = ache eimiedere on ciaiala mia lee en eneycra*y cielo ols nicl 52 Sciurus sp. (large red GAR) goocatospao eg oonuubonodoD por bodEco robo oon SODAD 115 Sastiguiniins MGA ogsacsnosacondsoooStouuedacpbT OOO DIR 2u carr a CRD Eb SUOOGNG Cogs 79 Shepherd dog (not a pure blood), FEU HLS ONE eect telsnetstetatone te ole ater= rodeo fenebatars lei ic eres 139 Rae ates hr Cay Mayan -sicters tejatater al eveaatevale ehnke rere 141 Sth, Same, (ORME) Soooeeodscse0ecboGboschor Oro CU rR GaR Aon SS Sp bcoSos 110 Solenodon paradoxus (young) = ---. 2-22... 22 ee ee ie ee ee i ee ieee 93 (YIITIN) ponédcodsecsod Od vaoonoe co OsOUGoUoRHC OGD CODD UROOor 3 938 INDEX vou. 35 PAGE Memur of Sorex (Shrew) ac cence yest eielolals else o cateie) ofetelslete a lolebelefal=tn terete ei=[ateta]tetel- ts t-toi- tele! otelel= 92 Shih Map BBBnUObO Ose Cobodearensrdoman 50 Saondonsao bond AGodD cums aoOboSCodonsaOC 141 Spelerpes ruber (red salamander) ......-.-- 2-22-21 e eee eee eee ee eee ee neee 43 Sphemodon™ punctate seperate tet etete tenants veka te ketene ete eee et ete 48 SHaridantey (Saas) soaoadapesgasog so ood0 7d 2UOSDdDOO Cog NHIsOusdOCHooagacengESSoSE 68 Sturnira’Vilium 3202 5<.c. ace tos See rere COE EO OE EOE Lee cle eee ec ereerr ne 76 Shitsh ((slassrlesjaton ints) opopedaonadnn negeaes064ADOdoo DO oNHONADOMOADadOIbOdowOUCON NOS 104 LO ove he, pile] Noy De oe amomooau od odoos Jedoe cous RUSS Gob come IOOS eat Sod a80 104 Tamandua tetradactyla ((amt-Cater ye ce crate teretees cuala ate aaa aes avariey colr oe teat Serena 112 TaphozoOus DHiLippiD NSIS) eeeer- yer inal deretene osetia helene kek rete etek tetsiete ened ote sate ceere noes ote 87 “UHL (KEV) oonon coocucnAcocse soo domgaUesnosSsehgnesase UDB ds soopp ec Setanconys 108 AMAA soba (name eblllon sat oaansensonogcancad soadc0ogOconcdoDCedg Nobo UOUS 112 TAUTOLTFAS US HOLY Re ctl Rvs tove cl otcicusts. Werersin are vere eReieteueset stn sleheote ueleiete elton se araeten repens torte tetettenete 107 Taxidea americana «(AMON Ca Dad SCI) eerie tete ratetcha tele exter arene ater nae eee 145 Testudo: ((Gopherus)) = DOlyp Mens. -wary-tereiete erchotertee eet shee etd oneee eked eee ate eae eee 59 Thylacinus eynNocephalus | (Masmayiane wiOlis) i rcnentetetetteretet ebarate eet terata lake iet atten er nena 119 Traculus jJavanicusn Gavia mM Ouse-d COL) me ryrrattstac islets ielalsratei tells tite etree eee 98 Trichosurus vulpecula (phalanger).................-- PRaconenaaobsesoGsLesdearS 6s 97 Trionyxspiniter ((soft-shelled tumtle)) ste cvesereteusy=tertetetete rele ete eae etanstetierenedetotiennten tara er 57 Tupaia. (tree Shrew)! ca, dsscce sveseuchays te et veps a eutoceas. © reaches pine gens ocr ten ett aeaereberenctetre 90 Turdus ‘migratorius (robin)! 2 [toec:.Sctesieaic.s ole ois Peas ot sles says fenel ay toe teterntey oteustone sie peredsteestobets 63 ‘LyMpanuchus) americanus s C(praLlencHiCKeM)) yan eisai etter tereteiieleietteretstetatel eters 61 Ursus americanus: (black: jbear))! se cpskerelovs sis de sete or mene sete needle cred eet tenet ee acer renee 106 maritimus (Polar Dear) meee lve re reel miele te Tarene Teyana arse terete raee aise Sieroter 1338 Vampyrops: Timeatusy 5 ots cticccveves.ccl< oe eyes aie voiecle vanes hsicbicteemeretec easiest lee toh ee ke TCL none 85 VAPAaNUS ANCMAR TUS) siciece ce seiec coe sessile tanct che yover atekeupucuane: stot: sve: shceckedeyetehs feet usp eactete aA ic ov esate ta vane 51 nuchalis(MOnitOL)) 2d ac cacisge sche oi dclersiale Vorier seisisioacisieve steel eastenere tires 51 SALW ATON : Se ey cvace cevsravensnegenshensRaevs love toresoney gle aietsVeseno sane clic tstekatonetate Cetencte rete act vate Reno rars 50 Vespertilio murinus:. Northwestern: (Umidvec occas iic sie cient te 190 167; Med. Dept: Northwestern s Unive. ccc crt ett ici teen 191 172, Med: Dept: Northwestern! Umive...c os serie cee cere ein 191 242, Cry Meds Goll ie coi sie cs ore bate oe tesa Ate edt tale te ciees cae eerie 191 145, Med. Dept. Northwestern Uniy........ pi ee ASS ae A 192 Ltd, Med. Dept. Northwestern Unive ciicrsceeie steele sietete hele tetetetere ete 192 15%, Med. Dept. North westenm (Uraiivecis sisi chvistreereiatcneeretenener 192 161, Med: Dept. Northwestern Unive sc scx cheercicic here eee emcees 193 153, Med; Dept. Northwestern’ Univis.es.eces ee ae ne tes 193 243. Cr: Meds" Coll.asace-iseen orci Cereee eee ae eee Me ee 193 148° Med. Dept. Northwestern Unive... esi. sei 194 Ve aa ) ae 215 On Of) | Reettemigaes naincaok oar concorde on os dot poo oducesp 194 97, Cris Med.’: Colles cy. ccrcuere ace sya cia exele oiele te olotensta stele is aka rchsiepetapareney acter 194 NO. 3 INDEX 239 PAGE EWAN. Oe Wyabive, jae INS GEE (Cine wit (hil Ge coogu Sanco roesosuGouodncasresbercaccone sk. 195 160 Med=sept., Northwesterns Unive sane nee eon eee ee ne oe 196 lib3 ye Med Dept. Nontiwesterni Unis eeee sees soe aan eee en: 196 Ibo veds Depts Northwestern! Unin aacenaeeate see tan eee 196 UG Vedas Dept NOGthiwestenns Univaae eee eect sane 197 bal MUCGE IDEs Wickless Whathia aan aqnens so scsossuesa4caness 197 LOO RCrse Ned {Colley cpenetrae crete ies ee een 198 150, Med. Dept. Northwestern Univ....:...............+....--.. 198 ioe Meds Dept. Northwesterm |Umives cess sneeer ene ce eeeee nae 199 1525) Med® Dept. Northwestern Umives..+....-0-25...2 9.05. bone. 199 Loe Weds Dept. Northwestern iUmives | eere aeeace oan nena 200 166) Meds Dept. Northwestern Unive--2 41.6. /20050 22 seen ee 200 (Suicide) Noel (ocr Med Coll weer een ase een 200 NOD MIS Or: SMed ai Collars ercjac net Coe wee yo ee 201 DIAC rowed SACOlle wrist caine ey ee eee Meat er ne 202 ANSTO ING) PEEP UWL Sh Nei Wiebe. soo donensAeab den somerenssarinloseconse. 202 VIRUS WEENIE, INS ES Ci, Wiech (Cail, Sab Ac ncn donb slondcs UohGeacutantech sane en. 202 147 Wed. eDeptmiN OCtwestenn iUMivasseiee sae eee ae een 203 (convict), No. 2, Med. Dept. Nebraska Wniv.......................... 204 No. 3h Wileelo IDET INAS, Winilizpconcdbonodcuoeseedssaucucuucuenc 204 2680 Croe Med si COlle as ricer Serrano ne oe ee ee 205 269 Cree MOG 4 COLE esp No y-sere eee erie eerie a eee ae 205 ZO MC rsa NCO iC Ollar iret eo Stee Skee vtec te eee ee 205 OTA CLES MECC sa GOMER Ea ee tery. te evi seciereore ee eat oS 206 Zee Orme NEU COM ae aa aenr eet An etna ae an ee ee 206 Lissa CE ee des COM taster erat ee Rie ih vse ee eee ee 206 PAs As ela yan O70) GUC eae reac aaa om Rete ae eee Wedel 207 PACs apd Oh ge ANY ICTEG KC Lo) DLE a ta vai pane, She ey Gi a eae eg a 207 216 Ore Wed WC OlltS eke nye cratarervie hetero ails thee a ie 207 ais: CreiMed Collie Senne Aster Aa saan oe tines Hea dee ee 208 ZS perp Medeg COMES cera venbrnciotintints Sy not eae eres 208 AO OLN NEC Collette aya ae amore setae err Meee Ee ee ee 208 280 2Cr-e Med Cole rire cers Aan Sh en ee Ooo res 209 Zo Ore Meds Colle turnin: | satcdasie eG acs hae ee 209 Ow OTL COs COM trseisy< sar aiiare (ASTRO oie ve teatro ee 209 2 Om OTe OCMC ONL Be cytes castles severe CICS oie ee ae ae ee 210 PANG) NOL FING Lis Glo DELO Soe tie aac a aSeR Le ny AS Ea nS 6 Ro 210 ZOO MOL MCA N COMM se tee teat aimee Her eee ne Oe te 210 PASTIS Cr oes Kets lot Cfo} NI Aiea te leu eee i ne ea eee Miami LENS Ou) 211 288; NOTA NC Oee COLL Mice ett sto ee anes eee ee ee ee 211 PASTS ( Ourt iN Kets Bol CX) DS ie Sie ears Perth ors en iene Mk onal ier oop ie 211 20, Osp Orn Led se CONES! VAN Seer heh etoen rok hye aicce en ae ae ee 212 (idiopathie epilepsy) cecseeciste,s er ease nee ae ere ee ee 212 Fetal human femora and their further developmenty xt. mcm hasr erie ee a Tag ISulgule), Oe Wea), INOS PEHCL NUE SEIN bie INGER on oaeoR BO n Ane bna geome bobdubodenbasaenonese.o- 162 HigMTess sroupedeaccordine, totyMeS4-16- 1s y7 onset sae oe cher oes aanee eeeoes ea oes 2 Hin SHAE secondwiny, pescombinatlonue ee see c= serene ie onan sven ane Sane 18 HIE Gin! anlual aA ers COM ACOs Kon cond bonne san ad oapAsndAant Lonmoadodsooueeneeeeoueaues. 18 Mins eSeccOnd wand uhindstypercombinationaes--4 re). 404. eee eee ance oe ene oe ee 18 LMI: UT NOWD. “Geong obdea see Cougs on Onan ae DEM EE On eA Tren be ee eS e See rtoite AEE are eae 12; 21 PDOSIELONS ie Oligr nem etter: ek cer ey RT oh capaho Sat ara Me eh eee 13; 21 RAMEE OF DEOMARINES Oi? WAS thpdeSoaodesbueebaos bop toceee sous osenesdceddeseuuesecu dec 17 HMC HOD Cem OlmOnm DOR ety ayo ttyl ae ae nike eee ee ee eee 26 (CHDUERE, OE Bite o.ag Gonerd 500 OUR ODER BORO ROS CeO ie AT en cr ERR eRe ae re Ad fe ae : 75 Generalscnaractervo une femunsoh amphibians). ea SoU QoSE pT DO amd 0.5 222, 223 | ee en Pena Sear mocdoUbdni outros eseinho co boDT Oo MSho se ac 224 \ 1 Pa eee Cee sate Tae cine cancobooncp Ocot oud sede vcs odcaenebouriss 225 (lf et ie eee ene ES Sranitciotid DOM Oe EIU COSC OSS ao TNE NEE OG Bt 226 ATA ed eee in OneNso Une or acs DOOODoD 06 Eo osomogurys Domo. 227 i Ga ee an Ee ean eo cAron oo pana cDOMD OTe eUe cas peat scedy Nor 228 D-H ananoacaooUUGNed Topco mDUaEatS oO Soe snap SHoacus 229 D6 (ee eR a bare eee een ae nA OAC Oe Hom ON OC TORO SOL SOL, 230 Table of bone types and lamellar divisions .............. GODS MCEO OTD ODORS TOMee adn cas 17 shapes in percentages......-2-- 2-51. -e eee eect eee ee eee ee centre eee 4 medullamy, indices) © oy... c.:):c irs tole mye otatbeelel «ile ete yece ete efaaslenehetcsslesens ohfehe teh egies tea iete 6 type combinations .........0. 60. s eee eee ee ee eee ee eee een eee 18 the distribution of the three bone typeS .........-..-2 +2222 eee eee eee eee eee 17 Third conclusion of the survey of averages of medullary HO GICES. shucks, ae cra sp antares Seo at Third type Of DONE ......... 6. cece cece rece e ree eee eee e eee e ee tenes terete rete etter ete et nee 14 tHOK StaSes OL’ Ae ciee Dee ee ot Oe EE Oe ee eee eres 15 Threefold division of lamelle. ... 2.2.56... c eee eee ee ee ee ee teens tees e eee 11 Tibia of negro, No. 248674, U. S. Nat. Mus. .......-.-.- 2.2 e eee eee ee ee ee eee eee 162 Total: number, Of Cemora ExaAMINed sam = series eo sched eke oe aka e tote what stat =yetet ote) ot= into fai eg tea eer 1 GA cH (cchll ee ee ee I niger cca 4 SmaeOrmadida SIO O OOS TATA DOSS meso 9 Ghntll ahhacitoe Mriche’ ganar osncosadcoomacno cb coouddeegooudacr dodo sda de oeSseeseasosss 11 Type advancement in amphibians ... Sibi oleh sate Oe See Eos RTA eee ny ots BETO TEs FIs ht TORR CL 18, 29 TODEU SSNs eae scose Pacers ns ca ctabete feimeas sl ele, Heke ened ape eh oYoaais eet) xv ne Cabs Sots teats eke te ees 19, 29 1th bys (ee Ae een Peet an er mre Ta an omic Ga e ac OS oo 19, 29 [ct |: ee ee RE Senev ety SUE os ir ria rath fence ate meet acurce to Ch8 ats 20, 29 pea shethiG: (rr re eon Rea rier A AA Rice OO ana ONG Se aD Oo, 20, 29 Teh mnnedoban abo steht cane ou Heo MERU Lr doo rear ication oS ona soso aus 21, 29 combinations eee erste eee eee eee sala tt cathe etre CR ee nee 18 1 ee ete a Ae Re ae cnr ss ROTO men DI GANS OAM ORIG.S CHU Co Oa toaina 18 TSDD De PS eS 8 Se dd anctave vetoes ake e Geass ree oe ET ote ee ois etter ey nce ge nee a near Be 18 A ht re rie oie eee mnie Soman aNten Maaco tacts 18 i FDAUDGy Weahaneanccn «cincango scope badbo nh scare cCaoo ue te jae terete sere 18 StEUCLUEG ACCOLGINE tO ClASSES OL AMINA SMe orretea elena eter cre apolar et ase gtet eae oa = tenet aa eer eae 18 Types) OL DONE: SCHUCHUTE He Ne wwicevetee vender talc eka etouch clas oF oilahey ste telsfe/lot Acteiciole els gol et ook en tee tet ee te Tee 12 Wina (Of NEETOWNO. 248674, Us Soe Nats MUSE. ere nts rele eieteretereceedet whetenas ketal nt eta lettatei toll of Right femur of Left femur of Right femur of Right femur of Right femur of Right femur of Right femur of Right femur of Cheiromeley Noctilio. Pteropus Pteropus Pteropus Pteropus Pteropus Pteropus Pteropus torquatus molossinus molossinus aldabrensis (Celebes) (Java) lepidus (Tongatabu) (small) - (large) FIFTY GENERA OF BATS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 9 Fic. 167 Fic. 168 Fig. 169 Fic. 170 Fic. 171 Fic. 172 Right femur of Left femur of Right femur of Echidna (egg-laying Left femur of Ornithorhynchus. Left femur of Left femur of Pteropus Tupaia. Tree mammal) (duckbill, egg-laying mammal) Cavia cutleri Scalopus aquaticus poliocephalus shrew (guinea-pig) (mole) (fruit-eating bat) Fie. 173 Fic. 174 Fig. 175 Fig. 176 Fic. 177 Fig. 178 Right femur of Right femur of Left femur of Right femur of Solenodon Femur of Lemur mongoz Left femur of Colobus Sorex (shrew) Macropus (wallaby) Solenodon paradoxus paradoxus (adult) abyssinicus caudatus (young) (African monkey) Fig. 179 Pic. 180 Fic. 181 “Fie, 182 Femur of Putorius vulgaris (weasel) Right femur of Mus rattus (black rat) Left femur of Heteromys Right femur of Myogale moschata (Spiny pocket rat) (desman) Fig. 183 Fig. 184 Irie. 185 Fic. 186 Femur of Cynomys ludovicianus Left femur of Trichosurus vulpecula Left femur of Phascolomys ursinus Right femur of Lasiopyga (prairie dog) (phalanger) (wombat) kolbi MAMMALS : SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 10 Fic. 187 Fia. 188 Fie. 189 Fic. 190 Fig. 191 Right femur of Tragulus Right femur of Mus Left femur of Erinaceus Right femur of Viverra Right femur of Ratufa javanicus (mouse-deer) sylvaticus (wood mouse) europezus (hedgehog) Civet maxima (giant squirrel) Fic. 192 Fic. 193 x Fic. 194 ; Fie. 195 Femur of Galeopithecus Left femur of Manis (scaly ant-eater) Right femur of Procayia Left femur of Helictis (flying lemur) capensis (coney) orientalis (asiatie badger) Tic. 196 Fic. 197 Fic. 198 Right femur of Cynocephalus (baboon) Right femur of Cynocephalus maimon Right femur of Hydrochwrus capybara ‘ (mandrill) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 11 Fic. 199 2 Fic. 201 Right femur of a fetal sheep (eleven weeks) Right femur of a fetal calf (eighteen weeks) Right femur of a fetal a (eight and one-half weeks) Fic. 202 Fia. 203 Fic. 204 Femur of Cariacus macrotis (deer) Right femur of Sus (domestic pig) Right femur of Sus scrofa (wild boar) Fra. 205 Fic. 206 Fia. 207 Fie, 208 , Femur of Alces machlis (elk) Right femur of Camelus (camel) Right femur of Auchenia glama Right femur of Rangifer (reindeer) (llama) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 12 Fic. 210 Fic. 211 Fic. 212 Right femur of Taurotragus Left femur of Connochztes Right femur of Ovibos (eland) taurinus albojubatus (gnu) moschatus wardi (musk ox) Fie. 209 Fie. 214 Fig. 215 Right femur of Tapirus (tapir) Right femur of Equus hemio- nus (wild ass of Asia) Fig. 218 Right femur of Potos caudiyol- vulus (kinkajou) Fig. 219 i Right femur of Lutra canaden- Right femur of Elephas africanus Fie. 217 ght femu = (Ateean Bleshant) Femur of Cholepus didactylus (two-toed sloth) sis (otter) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 13 Fig. 220 , Fig. 221 Fic. 222 Right femur of Simia satyrus (orang-utan) Left femur of Felis tigris Right femur of Hemigalus (tiger) hardwickii ‘ Fria. 223 Fie. 224 Fig. 225 Right femur of Tatu novemcinctus. Armadillo Right femur of Tamandua Left femur of Gorilla (gorilla) tetradactyla (ant-eater) Fig. 226 Fic. 227 Fic. 228 Femur of Presbytis rubicunda (monkey) Right femur of Hylobates (gibbon) Left femur of Anthropopithe- cus troglodytes (chimpanzee) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 14 Fie, 229 Fic. 230 Fig. 231 Right femur of Macacus rhesus (Indian monkey) Right femur of Sciurus sp. (large red squirrel) Right femur of Felis (domestie cat) Fig. 232 Fig. 233 Fic. 234 Via. 235 Left femur of Felis catus (wildcat) Iemur of Mephitis mephitica (skunk) Femur of Putorius vison (mink) Left femur of Cryptoprocta ferox (eatlike civet) Fig. 236 ic. 237 Fig, 235 Fig. 239 Right femur of Hyena crocuta (Hyena) Right femur of Thylacinus cynocephalus Right femur of Dasyprocta Left femur of Lasiopyga (Tasmanian wolf) i agouti eentralis johnstoni (monkey) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 15 Fic. 240 Fig. 241 Fic. 242 Right femur of Felis canadensis (Canada lynx) Right femur of Lasiopyga sp. (African Right femur of Midas rufoniger (South monkey) American monkey) lic. 243 Iria. 244 Fig. 245 Fic. 246 Left femur of Lemur variegatus Right femur of Lemur catta Left femur of Ateles (spider- Right femur of (ring-tailed lemur) monkey) (Tehuantepec) Callicebus torquatus (squirrel-monkey) ‘ Fic. 247 Fig. 248 Fic. 249 Left femur of Genetta (genet) Left femur of Pedetes (jumping hare) Right femur of Bradypus tridactylus (three-toed sloth) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 16 Fie. 250 Right femur of Castor canadensis (beaver) Fic. 253 Fic. 254 Fig. 255 Left femur of Equus caballus (horse) Right femur of Ovis (sheep) Right femur of Bison ameri- ecanus (bison) Fic. 256 Fic. 257 Fic. 258 Fig. 259 Right femur of a mule. Left femur of a mule. No. 229 Left femur of a mule. No, 235, Left femur of a mule. No. 236, No. 227, C. M. C. Cc. M. C. C. M. C. i Cc. MC. MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 17 Fig. 260 Fic. 261 Fic. 262 Left femur of Elephas indicus (Asiatic Right femur of Hippopotamus amphibius elephant) (hippopotamus) Fic. 263 Fic. 264 Right femur of Equus burchelli (zebra) Fic. 266 Fic. 267 Left femur of Ursus maritimus (polar bear) Right femur of Bubalis jacksoni (Hartebeest) Left femur of Phacocheerus (wart-hog) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 18 4 Fic, 268 Fic. 269 Fic. 270 Left femur of Felis concolor (panther) Left femur of Gulo luscus Left femur of Erignathus barbatus (seal) (wolverene) Fic. 271 Fic. 272 Fic. 273 Left femur of Bos bubalis (water buffalo) Left femur of Ovis montana (mountain sheep) Femur of Cephalophus (African antelope) Fic. 274 Fic. 275 2 : Palit eee Femur of Raphiceros (steinbok) Left femur of Gazella granti (Grant’s gazelle) Left femur of Kobus ellipsiprymnus f : (water buck) MAMMALS 18 SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 19 Fic. 277 Fie. 278 Fic. 279 Fic. 27916 Right femur of Arctomys monax Right femur of Canis latrans Right femur of Capra (goat) Right femur of a bull-dog (not a (woodchuck) (coyote) pure blood) Fic. 280 Fic. 280% Fic. 281 Right femur of a dog Femur of a fox terrier (not a pure blood) Fic. 282 Fic. 283 Fig. 284 Right femur of a mongrel dog Right femur of a bull dog (not a pure blood) Right femur of a collie dog (not a pure blocd) Fic. 285 Fie. 286 Right femur of a spaniel (not a pure blood) Left femur of Lepus cuniculus (rabbit) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 20 Fic. 287 Fic. 288 Tic. 289 Fic. 290 Right femur of Procyon lotor (raccoon) Os penis of Raccoon Femur of Canis lupus Right femur of a Felis leo (lion) (grey wolf) Fig. 291 Fia, 292 Right femur of Canis (small grey fox) Left femur of Taxidea americana (American badger) Fie. 295 Fic. 296 Fic. 294 Fic. 297 Fic. 298 Left femur of Canis aureus (jackal) Right femur of Haplondontia olympica, Left femur of a Erethizon (porcupine) sewellel (mountain beaver) MAMMALS SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 21 WM \ Oye ahh she ll SKC Bin 24 5 Fic. 299 Iie. 300 Fic. 301 Fig. 302 Right femur of a white Right femur of a white fetus Right femur of a white fetus of 4 months Right femur of a white fetus of 5-7 months fetus of 2-244 months of 3-3% months Fia. 303 Fic. 304 Fie. 305 Right femur of a white fetus of 8-9 months Right femur of negro fetus of 9 months Right femur of white fetus of 844 months (Craniorrhachischisis) ‘ ; Fic. 308 Left femur of a negro: No. 228481, U. S. N. M. Femur of mixed negro a Left femur of a negro. No. 3, Med. Dept., No. 247368, U. S. N. M. Tulane Univ. MAN (WHITE, BLACK) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL, 35, NO. 3, PL. 22 Fia. 309 Vie. 310 Fig. 311 Left femur of a negro. No. 87, M. D. T. U. Right femur of a negro. No. 7, M. D. T. U. Left femur of a negro. No. 4, M. D. T. U. Wie. 318 Fie. 314 Fic. 312 Right femur of a negro. No. 84, M. D. T. U. Left femur of a negro. No. 10, M. D. 'T. U. Right femur of a negress, age 40, No. 123, MDT As Fie, 315 Fic. 316 Fic. 317 Owed Left femur of a negro. No. 79, M. D. T. U. Left femur of a negro. No, 224714, U. S. N. M. Left femur of a negro. No. 11, M. D. T. U- MAN (BLACK) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 23 Fig. 319 Fie. 320 Left femur of a negro. No. 56, M. D. T. U. Left femur of a negress. No. 220, C. M. C. Fic Fic. 322 Fic. 325 Right femur of a negress. No. 220, C, M. C. Left femur of a negress age 14, Femur of a negro. No. 1, M. D. T. U. Amputated at lower third mixed black and white. No. 226, C. M. C. Fig. 324 Fie. 325 Right femur of a Kaffir negro. No. 263196, U. S, N. M. Right femur of a negro. No, 248674, U. S. N. M. MAN (BLACK) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 24 Fie. 326 Pie, 327 Fia. 328 Left femur of a negro. No. 248674, U. S. N. M. Tibia of a negro. No, 248674, U. S. N. M. Fibula of a negro. No. 248674, U. S. N. M. Fic, 329 Fic. 330 Ulna of a negro. No. 248674, U. S. N. M. Radius of a negro. No. 248674, U. S. N. M. Fic. 332 negro. No. 248674, U. S. N. M. Metatarsal bone of great toe of negro. No. 248674, U. S. N. M. Clavicle of a MAN (BLACK) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 25 Fig. 335 Fig. 336 Fic. 337 Right femur of a negro. No. 83, M. D. T. U. Right femur of a negro. No. 6, M. D. T. U. Fig. 338 Fic. 339 Fie. 340 Left femur of a negro. No. 5, M. D. T. U. Right femur of a negro. No. 8, M. D. T. U. Left femur of a negro. No. 7, M. D. T. U. rink oe a k __ Fig, 341 Fia. 342 Fig. 34244 Right femur of Pueblo Indian child one year Left femur of Pueblo Indian child six years old. Left femur of Pueblo Indian youth. No. old. No, 258675(z) U. S. N. M. No. 258675(L), U. S. N. M. 258675(S2), U. S. N. M. NEGRO. PUEBLO INDIAN. SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 26 Fic. 343 Fic. 344 Femur of Pueblo Indian adult. No. 258675(x) U. S. N. M. Right femur of Pueblo Indian adult. No. 227339, U. S. N. M. n Y. M. Left femur of Peruvian Indian. No. 266469(a), U. S. N. M. Left femur of Chicama Indian of Peru. No. 2, U. S. N. M. Left femur of Peruvian Indian. ‘0. 266469(b), U. Fia, 348 Fig. 349 I Right femur of Chicama Indian of Peru. Right femur of Chicama Indian of Peru. Left fimur of Chi Indian of Peru. No. 3, U. S. N. M. No. 1, U. S. N. M. No. 4, L N. M. MAN (YELLOW-BROWN) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 27 Fic. 351 Fic. 357 Fic. 360 Fig. 361 Left femur of Chicama Indian of Left femur of Pachacamac Indian of Right femur of Pachacamae Indian Left femur of Pachacamac Indian of Peru. No. 7, U.S. N. M. Peru (child). No. 12, U. S. N. M. of Peru (adult). No. 15, U. S. N. M. Peru. No. 7, U. S. N. M. Fie. 362 Fic. 363 Fic. 363a Right femur of a Japanese male. No. 245, Femur of Egyptian child of XII Dynasty. No. 256479(de) Vascular origin of an Haversian Cc. M. C: U. S. N. M. system as seen in Fig. 363 at A : f ; Pia. 364 d i Fig. 365 Fic. 366 Femur of Egyptian child of XII Dynasty. No. 256479 (d) Right femur of Egyptian child of XII Dynasty. Femur of Egyptian youth of XII Dynasty. U.S. N. M. No. 256479(a3), U. S. N. M. No. 258675(a) U. S. N. M. MAN (PERUVIAN INDIAN AND EGYPTIAN) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 28 Fic. 367 Fig. 368 Fie. 369 Right femur of Egyptian adult -of XII Dynasty. Left femur of Egyptian adult of XII Dynasty. Right femur of Egyptian adult, XII Dynasty. No. 256481(d), U. S. N. M. No. 256481(a), U. S. N. M. ' No. 258675(e), U. S. N. M. us mile 9) Fig. 370 Fic. 371 Femur of Egyptian adult of XII Dynasty. Right femur of Egyptian adult of XII Dynasty. No. 256478(23), U. S. N. M. No. 256478(x) U. S. N. M. Hie. 372 . : Fie. 373 Femur of a male white. No. 1629, U. S. N. M. Right femur of a female white. No. 147, M. D. N. U. MAN (EGYPTIAN AND WHITE) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 29 Fia. 374 Fie. 375 Fic. 376 Right femur of a white child less than one year old. Femur of a male white. No. 53, C. M. C. Right femur of a male white. No. 171, M. D. N. U. No. 249588, U. S. N. M. Fia. 377 Fic. 378 Fic. 379 Left femur of a male white. No. 95, C. M. C. Left femur of a male white. No. 96, C. M. C. Left femur of a male white, age 45. No. 168, M. D. N. U. 3 Fig. 381 Fig. 382 Left femur of a male white, age 50. No. 10, Iemur of a female white, age 52. No. 227876, Femur of a female white, age 60, No. 227880, M. D. N. U. U.S. N. M. U.S. N. M. MAN (WHITE RACE) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 30 Fia. 384 Fia. 385 Fig. 386 Fig. 387 Left femur of a male white. Right femur of a male white. Right femur of East Indian male. Left femur of No. 223 C. M. C. No. 162, M. D. N. U. No. 244, C. M. C. No, 223, C. M. C. amputated Fig. 388 Fig. 389 Fie. 390 Right femur of a male white. No, 228479, Right femur of a male white, age 45. No. 154, Left femur of a male white. No. 146, . S. N. M. M. D.N. U. M. D. N. U. Fig. 391 Fie, 392 ia, 393 Left femur of a male white. No. 159, Right femur of a male white. No. 167, Right femur of a male white. No. 172, M. D. N. U. M. D. N. U. M. D. N. U. MAN (WHITE RACE) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 31 Fie. 394 Fic. 395 Fic. 396 Right femur of a male white. No. 242, Right femur of a male ite, age 60. No. 145 Left femur of a female white. No. 174, Cc. M. C. M. D. N. UL M. D. N. U. J Fig. 397 Fig. 398 Fig. 399 Right femur of a male white. No. 157, Left femur of a male white. No. 161, Right femur of a male white. No, 153, M. D. N. Uz M. D. N. U. M. D. N. U. é Fie. 400 Fic. 401 Fig. 402 Right femur of a male white, No. 2438, Left femur of a male white. No. 148, Left femur of a male whit>. No. 230, Cc. M. C. M. D. N. U. Goat G: MAN (WHITE RACE) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 32 Fic. 403 Fic. 404 Fis, 4 Left femur of a male white. No. 97, C. M. C. Left femur of a male white. No. 99, C. M. C. Right femur of a male white. No. 160, M. D. N. U. Tic. 406 Fic. 407 Fic. 408 Left femur of a male white. No, 163, M. D, N. U. Right femur of a male white. No. 156, M. D. N. U. Left femur of a male white. No. 169, M. D. N. U. Fie. 409 Fic. 410 Fig. 411 Right femur of a male white, age 35. No. 151, Left femur of a male white. No. 100, C. M. C. Right femur of a female white. No. 150 M. D. N. U. M. D. N. U. MAN (WHITE RACE) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 33 Fic. 412 Fie. 413 Left femur of a male white. New bone development seen on the right. Left femur, 33 mm. below the section 412. The femur has become No. 152, M. D. N. U. double. No. 152, M. D. N. U. Fic. 414 Fic. 415 Fie. 416 Left femur of a female white. No. 164, M. D. N. U. Left femur of a female white. No. 166, M. D. N. U. Right femur of a male white, age 22 (suicide). No. 175 Fic. 417 Fie. 418 Left femur of a male white. No. 98, C. M. C. Left femur of a male white. No. 91, C. M. C. MAN (WHITE RACE) SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 34 Fig. 419 Tic. 420 Fic, 421 Femur of an Australian. No. 227420, U. S. N. M. Left femur of a male white. No. 94, C. M. C. Right femur of a male white. No. 142, M. D. N. U. MAN (AUSTRALIAN; WHITE RACE) Fie. 423 Fic. 424 Fic. 425 Fie. 426 A single Haversian system Haversian system showing Haversian system showing Haversian system showing enlarged showing lamelle early stage of senility later stage of senility latest stage of senility DIAGRAMS SHOWING STAGES OF SENILITY SMITHSONIAN CONTRIBUTIONS TO KNOWLEDGE VOL. 35, NO. 3, PL. 35 Fic, 428 ; Fie. 429 Fic. 436 Right femur of male white (convict). No. 2, Right femur of a male white (convict). No. 3, Right femur of a male white. No. 274, C. M. C. M. D. Neb. U. M. D. Neb. U. Fic. 437 Vic. 439 Fic. 445 Right femur of a male white. No. 275, Right femur of a male white. No. 277 C. Mf. C. Cc. M. C. Fia. 451 Fie, 452 Fie. 453 Right femur of a male white. No. 289, Left femur of a male white. No. 296, Left femur of a male white, age 40 (case of Cc. M. C. Cc. M. C. idiopathic epilepsy). No. 1, N. S. H. MAN (WHITE RACE) LG! ei , : ei | 2 _ Cae ; i 5 i ; | ey ) yay eee ae. 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