CATS ct PASSE Ane _ so i iV ? % * ee y Lie ad e” . te - = eh y ay SMITHSONTAN MISCELLANEOUS COLLECTIONS VOL.93 “EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES, AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN ”"_sMITHSON (PUBLICATION 3340) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1935 The Lord Waltimore Press BALTIMORE, MD., U. 8. As t ve we ADVERTISEMENT The present series, entitled “ Smithsonian Miscellaneous Collec- tions,’ is intended to embrace all the octavo publications of the Institution, except the Annual Report. Its scope is not limited, and the volumes thus far issued relate to nearly every branch of science. Among these various subjects zoology, bibliography, geology, mineralogy, anthropology, and astrophysics have predominated. The Institution also publishes a quarto series entitled ‘ Smith- sonian Contributions to Knowledge.” It consists of memoirs based - on extended original investigations, which have resulted in important additions to knowledge. C. G. Assort, Secretary of the Smithsonian Institution. (ii) bo N Q. CONTENTS Wricut, W. H., and McAtister, E. D. The effect of ultraviolet radiation on the ova of the ascarid roundworms Toxocara canis and To-xascaris leonina. 13 pp., Dec. 26, 1934. (Publ. 3201). Scumitt, Watpo L. Mud shrimps of the Atlantic coast of North America. . 21 pp., 4 pls., Feb. 15, 1935. (Publ. 3292.) GaTEs, G. E. New earthworms from China, with notes on the synonymy of some Chinese species of Drawida and Pheretima. FOr pps, 15 figs., Feb: 27, 1945. (Publ. 2202.) RANDERS-PEHRSON, N. H. Pioneer wind tunnels. 20 pp., 4 pls., 8 figs., Jan. 19, 1935. (Publ. 3294.) RessER, CHARLES ELMER. Nomenclature of some Cambrian trilobites. 46’pp., Feb: 14, 1935. (Publ. 3295.) HRplLicKa, ALES. Ear exostoses. 100 pp., 5 pls., May 14, 1935. (Publ. 3296.) McAttster, E. D. The Christiansen light filter: its advantages and limitations. 12 pp., 2 pls., 4 figs., Apr. 2, 1935. (Publ. 3297.) Basser, R. S. The classification of the Edrioasteroidea. 11 pp., E pl, Apr: 4, 1935. (Publ. 3307.) CANU, FERDINAND, and BAssLer, Ray S. New species of Ter- tiary cheilostome bryozoa from Victoria, Australia. 54 pp., © pls., Apr: 26, 1935. (Publy 3302) STRONG, WILLIAM DuNCAN. An introduction to Nebraska ar- cheology. 323 pp., 25 pls., 30 figs., July 20, 1935. (Publ. 3303.) (v) -_ mee <4)! >. ie ee ‘aly oh i 7" +a s eel bo? => = sy ae y? a i a ie | ® ea . At a ae “2 _ a > es A Pot SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 1 THE EFFECT OF ULTRAVIOLET RADIATION ON THE OVA OF THE ASCARID ROUNDWORMS TOXOCARA CANIS AND TOXASCARIS LEONINA BY W. H. WRIGHT Zoological Division, Bureau of Animal Industry, U. S. Department of Agriculture AND E.. D. MCALISTER Division of Radiation and Organisms, Smithsonian Institution (PUBLICATION 3291) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION DECEMBER 26, 1934 The Lord Baltimore Press BALTIMORE, MD., U. S. A. Hie BEPECM OF ULTRAVIOLET, RADIATION ON THE OVA OF THE ASCARID ROUNDWORMS TOXOCARA CANIS AND TOXASCARIS LEONINA By W. o. WRIGHT, Zoological Division, Bureau of Animal Industry, U.S. Department of Agriculture AND E. D: McALISTER, Division of Radiation and Organisms, Smithsonian Institution INTRODUCTION Through the courtesy of Dr. C. G. Abbot, Secretary of the Smith- sonian Institution, the opportunity was presented to study the action of ultraviolet light on the ova of the two species of ascarids commonly infesting the dog. It was desired in the experiments reported here to determine, if possible, the radiotoxic effect of ultraviolet light of different wave lengths and, as a practical measure, to correlate the possible sterilizing action of sunlight on the ova of these two species of ascarids. The apparatus employed for the radiation of the ova, described in detail by Brackett and McAlister (1932), consisted of a quartz mono- chromator and a quartz mercury arc. This apparatus eliminates the disturbing effect of the large amount of heat attending all total arc exposures and permits irradiation by the different wave lengths of the mercury spectrum. The intensities of the spectral lines employed were as follows: Wave length (A) Intensity (ergs/sec. cm”) 3650 9500 3130 4200 3022 1900 2967 gI0 2804 610 2052 570 In correlating the dosage employed with ultraviolet intensities in sunlight, the determinations of Coblentz and Stair (1931) have been used. These investigators gave the value of 65 microwatts per cm’ (650 ergs/sec. cm’) for noonday, midsummer sun at Washington, SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 1 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 D. C., for wave lengths of 3130 A to the solar limit at 2970 A. From their data there have been computed an average value of 55 microwatts per cm’ (550 ergs/sec. cm’) for the above-mentioned wave lengths in July sunlight between 9: 00 a. m. and 3: 00 p. m. at Washington, D. C. This value has been used in translating into terms of days the exposures employed in some of the tests reported here. The ova of Toxocara canis are globular, subglobular, or slightly ovoid in shape and of a light brown color. Externally, the egg is covered with a thick, albuminous coating which is mammillated at regular intervals. The eggs vary in measurement from 82 to 102 p in length and 79 to 96 » in width. The ova of Toxascaris leonina are ellipsoidal to subglobular or globular in shape, with thick, double- contoured, smooth, clear shells. They measure 82 to 96 » in length and 82 to 92 w in width. REVIEW OF LITERATURE The effect of ultraviolet irradiation on the ova of Ascaris equorum (= A. megalocephala) has been studied in some detail by a number of investigators. Most of these experiments were conducted from the standpoint of the cytologist with a view to determining the effect of the light on various parts of the egg or on the whole egg at different stages of development. Stevens (1909), using a total arc exposure, found that exposure of the whole egg to ultraviolet light for 6 to 8 hours did not usually kill the egg at once, but prevented further development when the eggs were in the 2- to 4-cell stage at the time of exposure. Exposure for a period too short to prevent further cleavage (4 to 3 hours) caused various irregularities in development including irregular fragmentation of the chromosomes, delay in cleavage and abnormal gastrulation. Stevens believed that ova arrested in develop- ment may be said to be paralyzed to such an extent that they are unable to initiate any further mitosis. In his extensive experiments in exposing various parts of the egg of A. equorum to ultraviolet light Schleip (1923) employed the micro- scopic method devised by Hertel (1904) and modified and improved by Tschachotin (1912). The apparatus used gave a spectral line of 2800 A from the magnesium arc. Schleip found that irradiation of the whole egg or various parts of the egg for varying periods of time resulted in a marked lethal effect. Schleip concluded, moreover, that no part of the egg could be influenced by ultraviolet light without producing secondarily some alteration in other parts not exposed to the light. Ruppert (1924) also employed the apparatus of Tschachotin in irradiating the ova of A. equorwm. In Ruppert’s experiments short NO. I IRRADIATED ASCARID OVA——-WRIGHT AND McALISTER 3 exposures of the whole egg resulted in marked abnormal embryonal development, which appeared most frequently in the gastrula stage. Longer exposures gave a marked lethal effect. Ruppert concluded that there is a rhythmic alteration in the lethal effect of ultraviolet light, depending on the stage of development of the egg at the time of exposure. Seide (1925), in irradiating ova of A. equorum, used the method of Tschachotin as well as total arc exposures at wave lengths between 4050 and 2530 A of the mercury spectrum. In his experiments there was no apparent lethal effect on the eggs and no apparent lag in development. Nolf (1932) observed that a very small total arc exposure at wave lengths between 2800 and 3150 A or between 1800 and 3150 A was sufficient to prevent a large percentage of the ova of Ascaris lum- bricoides from reaching embryonation. A slightly greater exposure was completely lethal to the eggs. Although results obtained by the above-named investigators are of general interest, their findings offer no means of comparison with results of experiments reported in this paper, in which measured in- tensities of single wave lengths of ultraviolet light were employed. The intensity of the 2800 A spectral line from the magnesium are was not reported by those using this method, making comparison impos- sible, and it is equally impossible to compare results with those of workers who employed total arc exposures. EXPERIMENTS SERIES A Preliminary tests were made by exposing Toxocara and Toxascaris eggs to six wave lengths of ultraviolet light in order to gain some idea as to lethal effect, if any, of these various wave lengths. The ova of both species of ascarids were mixed and placed on a glass slide and allowed to dry at room temperature. The slides were exposed to ultra- violet light for a length of time sufficient to provide an equivalent dosage (the product of time and intensity ) at the various wave lengths. As the longest period of exposure was 20 minutes, each slide was dried for 20 minutes, including the time of exposure to the light, in order to provide equal conditions for the test. Each slide was then ex- posed to the arc for the time stated in Table 1. The dosage was 684,- 000 ergs/cm’ or approximately equivalent for the 3022 A slide to 18 minutes exposure to nooday, midsummer sun at Washington, D. C. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 The exposed area on each slide was marked off with a diamond point ; this area was equivalent to the dimensions of the light ray dis- charged from the aperture, or 5 by 30 mm. Eggs on the same slide outside of the irradiated area served as controls ; the control eggs were therefore subjected to identically the same conditions during develop- ment as were the irradiated eggs. After exposure, each slide was immediately placed ina Petri dish and covered with a 1 per cent solution of formalin, in which the eggs were permitted to develop. For the most part the eggs adhered to the slide. The culture was allowed to develop for a period of 8 to 9 days at temperatures ranging from 26° to 28° C. At the end of this time, counts were made to determine the TABLE 1.—Results of Exposure of Ova of Toxocara canis and Toxascaris leonina to Ultraviolet Light Series A—Exposed on May 23, 1934 Dosage—684,000 ergs/cm? Percentage embryonated ee oO Ne. Wave exposure Dare Toxocara Toxascaris slide length count Minutes} Seconds frag Control ied Control A 1934 I 2652 20 =e May 31 27.5 32 6.5 25 2 2804 18 46 May 31 46 48 6 29.5 3 2967 12 30 June I 51 50.5 20 22 4 3022 6 if June I 60.5 57 33 2205 5 3130 2 44 June rt 49.5 47 27.5 23.5 6 3650 I 12 June I 47 49.5 26 27 percentage of embryonation in the eggs of the two species, both on the control part of the slide and on the irradiated part of the slide. For each count, 200 ova were taken. Table I summarizes the results obtained from the irradiation of ova in series A. From an examination of the data it is apparent that the relatively short exposure used in this series had little or no effect on the develop- ment of the ova in the majority of cases. However, in two instances the ultraviolet light appears to have exerted a definite toxic effect on the ova of Toxascaris. On slide 1 (wave length 2652 A) only 6.5 per cent of the irradiated ova became embryonated, whereas 25 per cent of the control ova became embryonated. On slide 2 (wave length 2804 A), 6 per cent of the irradiated ova were embryonated at the time the count was made, whereas 29.5 per cent of the control ova were INO Seek IRRADIATED ASCARID OVA—WRIGHT AND McALISTER 5 embryonated. In each of these instances it is apparent that the ultra- violet light prevented development in a relatively large proportion of the Toxascaris eggs. In other cases the differences in embryonation were well within the limits of experimental error in the counting technic. SERIES B In view of the fact that wave lengths from 2652 to 2967 A are below the limits of the solar spectrum, there appeared to be little information of practical value to be derived from the further irradiation of ascarid eggs at these wave lengths even though Toxascaris ova were consider- ably affected by irradiation at wave lengths of 2652 and 2804 A. For this reason further experiments were confined to irradiation of the ova at wave lengths within the range of the solar spectrum with a view to ascertaining the relative lethal effect of sunlight, exclusive of heat and desiccation, on the ova of these two species of ascarids. This point has practical application in the control of ascariasis. In series B the ova were exposed to a dosage approximately equal to 40 times that used in series A. The dose was equivalent to 27,400,000 ergs/cm’. This exposure for dish I (wave length 3022 A) was ap- proximately equivalent to 12 hours of noonday, midsummer sun at Washington, D. C. For this test, a mixed culture of Toxvocara and Toxascaris ova was dried in the bottom of 50-mm culture dishes for 10 minutes. After the eggs had dried on the bottom of the culture dish, water was added to the dish to a depth of 2 mm. This prevented drying of the culture during the period of irradiation. An area equivalent to the light aper- ture, or 5 by 30 mm, was marked off on the bottom of each culture dish with a diamond point and the eggs within that area were exposed to the ultraviolet light. Eggs without this area were not exposed and were used as controls. After exposure of the eggs, formalin was added to the culture dishes to provide a concentration of I per cent in order to prevent bacterial growth in the cultures. Temperatures during de- velopment of the cultures ranged between 29° and 30° C. Counts were made nine days after irradiation ; 200 ova were taken in each count. The results of the experiment are recorded in table 2. In only one case was there any apparent toxic effect from the ultra- violet irradiation in series B. In dish I (wave length 3022 A) there resulted a marked lethal effect on the ova of both Toxocara canis and Toxascaris leonina, although the effect was most marked on the ova of the latter species. Of the ova exposed at this wave length, 24.5 per cent of the irradiated Toxocara eggs developed to embryonation, where- 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 as 58.5 per cent of the control eggs became embryonated. Only 8 per cent of the irradiated Tovrascaris eggs developed, whereas 29 per cent of the control eggs became embryonated. A slight toxic effect may have been exerted on the ova exposed in dish 2 (wave length 3130 A), although reference to the series C experiment would seem to indicate that the differences noted above are probably due to chance variation in the counts. TABLE 2.—Results of Exposure of Ova of Toxocara canis and Toxascaris leonina to Ultraviolet Light Series B—Exposed on June 5, 1934 Dosage—27,400,000 ergs/cm?2 Percentage embryonated Bar fo) J - Date N . Wave EES ISSUERS of Toxocara Toxascaris dish length count Hours |Minutes a Control trad Control A 1934 3022 4 ae June 14 24.5 58.5 8.0 29.0 2 3130 I 57" | June 14 43-5 51.0 22.5 30.0 g 3650 A 48 June 14 56.5 51.0 28.5 30.0 8 7 percent overdose. SERIES C In series C irradiation of Toxocara and Toxascaris ova with ultra- violet light was prolonged to the time indicated in table 3. The dosage used was approximately five times that employed in series B, and amounted to 137,000,000 ergs/cm’. For dish 1 (wave length 3022 A) the exposure utilized was approximately equivalent to 60 hours of noonday, midsummer sun at Washington, D. C., or 12 days of July sunlight. The method of exposure used in series B was found unsatisfactory from the standpoint of making microscopic counts. Owing to the long exposure in series C, it was necessary to devise a more suitable method of irradiating the eggs so that drying would be prevented. There were constructed small glass dishes 5 mm wide, 30 mm long, and 5 mm high, the first two measurements representing the size of the area over which the light was dispersed from the aperture. A mixed culture of Toxo- cara and Toxascaris eggs was placed in the dishes and water added to a depth of approximately 3 mm. As evaporation proceeded during the course of irradiation, more water was added to prevent drying of the eggs. NO. I IRRADIATED ASCARID OVA—-WRIGHT AND McALISTER 7 A number of eggs sufficient to cover only the bottom of the dish was used in order to prevent any overlapping and shadowing of eggs from the ultraviolet light. After irradiation the eggs were transferred by means of a clean pipette to culture dishes containing a 1 per cent solution of formalin. As a control, a culture was made in 1 per cent formalin on the same date as the first irradiation exposure and was subjected to the same conditions of development as were the irradiated cultures. Temperatures during the period of development varied be- tween 30° and 35° C., as recorded on a thermograph chart. Table 3 gives the results of this experiment. In this series 400 ova were taken for each count. TABLE 3.—Results of Exposure of Ova of Toxocara canis and Toxascaris leonina to Ultraviolet Light Series C—Exposed July 16, 18 and 20, 1934 Dosage—1I 37,000,000 ergs/cm?2 Warstion Percentage embryonated No. fo} Wave Date Date a length esol IRS of of Toxocara Toxascaris exposure count : ! Hours |Minutes Irradiated | Control | Irradiated | Control A 1934 1934 Tn gO22 20 ste July 16 | July 27 20.5 65-5 1.5 42.0 2553130 9 10 | July 20 | July 31 63.5 65.5 29).7.5) || 42-0 3650 4 if July 18 | July 28 | 63.0 65.5 46.5 42.0 It will be noted that the exposure to ultraviolet light at a wave length _ of 3022 A was the only exposure which produced any apparent toxic effect on the eggs of either species. The counts indicated that only 20.5 per cent of the irradiated Toxocara ova reached embryonation as com- pared with 65.5 per cent of the eggs embryonated in the control culture, and only 1.5 per cent of the irradiated Toxascaris ova became em- bryonated as compared with 42 per cent of the eggs embryonated in the control culture. In this experiment the irradiated and control cultures were examined daily under the microscope. A careful check was made on the develop- ment of the cultures in an effort to determine whether there was any lag in development in those ova which actually started to segment. No such lag in development was noted. Those ova which segmented ap- parently began segmentation as promptly as did nonirradiated ova in the control culture. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 The radiotoxic effect on the ova of both species was exhibited in different ways. On the ova of both Torocara and Toxascaris it was apparent in some cases that the ultraviolet light had some direct and undelayed lethal effect, inasmuch as many eggs never underwent any cleavage. On the other hand, some of the ova of both species developed in part until development was definitely arrested. In order to check on this point, the culture in dish 1 (wave length 3022 A) was re- counted at the expiration of 21 days in an effort to determine whether any ova which were in intermediate stages of development at the time of the first count, 11 days after exposure, had completed their develop- ment. It was found that none of the ova which were partially developed at the time of the first count had completed their development at the time of the second count 10 days later. It would appear that such ova had been definitely and permanently arrested in development, a cir- cumstance which agrees with the results obtained by Stevens (1909), who believed that such ova may be said to be paralyzed to such an extent that they are unable to initiate any further mitosis. In some of the ova arrested in the course of development it was apparent that cleavage had proceeded normally up to a certain point. On the other hand, all sorts of irregularities were noted in the develop- mental stages of some of these eggs. These irregularities were most frequent in Toxocara ova. In some the cytoplasm was apparently degenerated and was distributed in various areas within the shell. In other ova the cytoplasm contained large vacuoles. Irregularities and abnormalities in blastulation and gastrulation were marked. Some of the Toxocara ova contained partly formed embryos, the unformed remainders of which were composed of undifferentiated and irregularly formed masses of cells. In some of the eggs the first somatic stem cell failed to develop, whereas the first germinal stem cell divided many times. In other eggs the first germinal stem cell developed only partly. In most of the Tosxascaris ova that showed development it would appear that segmentation proceeded normally up to the 8- to 16-cell stage, at which stage it was definitely arrested. Development in none of these ova proceeded to the blastula or gastrula stage, except in a very few eggs which actually reached embryonation. In all ova of both species which became embryonated, the larvae appear to have ensheathed normally. In order to determine the infectivity of embryos which developed in the irradiated ova, on August II, 1934, the three cultures irradiated in series C were each fed to a young albino rat; at the same time the control culture was fed to a rat from the same litter. The rat which received the control culture gave birth to a litter of 10 young on August IN(ORe gE IRRADIATED ASCARID OVA——-WRIGHT AND McALISTER 9 17, 1934. As it was desired to raise this litter, the animal was not killed. The three other rats were killed on August 17, 18, and 21, 1934, respectively. Ascarid larvae were recovered from the liver and lungs of each of the three animals. All of these larvae proved to be those of Toxocara canis. Failure to recover Toxascaris larvae from the rats was not unexpected, as only a very limited supply of To.xascaris leonina ova was available at the start of the experiment and the cultures con- tained only a relatively few ova of this species. Failure to recover larvae is therefore not regarded as definite evidence that the embryos within the ova were not infective. In all of the experiments in which any radiotoxic effect was noted from ultraviolet irradiation, Toxascaris eggs appeared to be the most severely affected by the exposure. In series C, the lethal effect ap- parently resulted in a definite killing of the ova without any segmen- tation, or in an arrest in development during the early stages of seg- mentation. In no case did Toxascaris eggs which were arrested in development reach the blastula or gastrula stages. This is in contrast to the effect on Toxocara ova in which blastulation or gastrulation, even though abnormal, was reached in some cases. It would appear that the more marked radiotoxicity on the eggs of Toxrascaris may be associated with the smooth, rather clear, nonmammillated shell of these eggs, in contradistinction to the mosaically formed, mammillated, more darkly pigmented shell of Toxocara. It is believed that the mosaic pattern of the shell of this species of egg would tend to diffuse the ultraviolet rays and that the deeper pigmentation of these eggs would be responsible for some absorption of the ultraviolet spectrum before the rays had had an opportunity to reach the cytoplasm or nucleus of the egg. CORRELATION OF PRESENT RESULTS WITH PREVIOUS INVESTIGATIONS A number of investigators have reported on the effect of exposing the ova of various species of ascarids to sunlight. Ross (1916) re- ported that the eggs of the human ascarid, Ascaris lumbricoides, de- veloped and remained alive when kept for 6 weeks on glass slides in the direct sunlight in India; Ross was of the opinion that a relatively high humidity was not essential for the development of the eggs. Ross’s observation is at variance with the results obtained by Mana- lang (1927), who found that human ascarid eggs on glass slides were all dead after 14 hours’ exposure to the direct sunlight in the Philip- pines, although such eggs resisted an exposure of one-half hour. How- ever, Ohba (1926) reported that ascarid eggs in water cultures in IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 direct sunlight continued to develop. Brown (1927) exposed the eggs of A. lumbricoides to direct sunlight in Panama by placing them in sand, and after a period of 21 days of such exposure it was found that all of the ova had degenerated. Brown considered that the destruction of the ova was due to two factors, viz, high temperature and desic- cation. Soil temperatures in the sand cultures were found to reach at least 123° F., which appears to be above the lethal range of heat for ova of this species. Caldwell and Caldwell (1928) exposed fecal-soil cultures of the ova of the human and pig ascarids to sunlight in Ala- bama, and after a period of 3 days’ exposure, the ova in all cultures were disintegrated. The maximum temperature recorded in the cul- tures was 146° F. These investigators place great emphasis on desiccation as the chief lethal factor involved, inasmuch as cultures moistened at hourly intervals while exposed to sunlight showed little disintegration of the ova of the pig ascarid ; ova of the human ascarid showed less resistance. Otto (1929) reported that eggs of the human ascarid developed and remained alive over the summer of 1928 on the surface of clay, loam, sand, and cinder-loam soils in the shade in south- western Virginia. Many of the eggs on the first three soils in the sun died rapidly, but after 160 days about one-fourth of the eggs isolated were still alive, whereas most of those on cinders in the sun died before becoming embryonated. The high temperatures recorded on the surface of the cinders lead the author to conclude that temperature played an important part in the death of these eggs. Apparently, Owen (1930) is the only investigator to observe the effect of sunlight on the ova of Toxocara canis. Owen exposed the ova of this species to summer sunlight in Kentucky and Minnesota and found that such ova disintegrated before reaching the infective stage. A surface-soil temperature of 131.9° F. was obtained on the plots of soil on which the eggs were exposed. Owen was of the opinion that the failure of the eggs to develop was due to the high temperature. Schwartz (1932) reported that ova of Ascaris vitulorum did not survive exposure on glass slides to 1 hour’s direct sunlight in the Philippines. Eggs exposed in beakers of water also failed to survive after 1 hour of direct sunlight. Schwartz then exposed ova in vials painted with India ink to exclude light. An exposure of I hour in sunlight was lethal to ova in painted and unpainted vials, and Schwartz thus attributed the death of the ova to the temperature, which reached asec, Ohba’s negative results from exposure of ascarid ova to sunlight are difficult to interpret in the light of our irradiation tests, as the length of exposure is not stated in the English summary of Ohba’s NO: I IRRADIATED ASCARID OVA—-WRIGHT AND McALISTER Mail paper. Regardless of these negative results, it is apparent from our tests that sunlight, through the ultraviolet spectrum at wave lengths of approximately 3022 A, does exert a definite radiotoxic effect on ascarid ova. However, owing to the relatively long exposures necessary for the development of this lethal effect, it appears probable from a practical standpoint that other factors, such as desiccation or high temperatures, exert a more destructive action on these ova. In humid, tropical climates, however, ultraviolet light probably does serve in some measure in preventing the development of such ascarid ova as are directly exposed to sunlight. SUMMARY AND CONCLUSIONS Irradiation of the ova of Toxocara canis and Toxascaris leonina at measured wave lengths from a quartz monochromator and quartz mercury arc resulted in a certain degree of radiotoxicity to the ova exposed at certain wave lengths. A dosage of 684,000 ergs/cm’* at wave lengths of 2652 and 2804 A had a marked lethal effect on the ova of Toxascaris leonina but ap- parently no effect on the ova of Toxocara canis. A similar dosage at wave lengths of 2967, 3022, 3130, and 3650 A was without effect on the ova of either species. A dosage of 27,400,000 ergs/cm’ at a wave length of 3022 A re- sulted in definite radiotoxicity on the ova of both species of ascarids. Exposures to the same dosage at wave lengths of 3130 and 3650A showed no effect. A dosage of 137,000,000 ergs/cm* at a wave length of 3022 A showed a marked lethal effect on the ova of both species. In the case of Toxocara, only 20.5 per cent of the irradiated ova developed to embryonation as compared with 65.5 per cent embryonation in the control culture; only 1.5 per cent of the Toxascaris ova reached embryonation as against 42.0 per cent embryonation in the control culture. The dosage employed was approximately equivalent to an exposure of 60 hours noonday, midsummer sun at Washington, D. C., or 12 days of average July sunlight. In all of these tests, the ova of Toxascaris leonina proved more susceptible to the action of ultraviolet light than did the ova of Toxo- cara canis. This difference is probably accounted for by the difference in structure and pigmentation of the shell; the mosaically patterned, mamumillated, darkly pigmented shell of To.ocara ova would appear to disperse and to absorb more light than does the clear, unmammillated, lightly pigmented shell of Tosxascaris ova. [2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Although the marked lethal effect of sunlight on ascarid ova, as reported by several workers, is probably due chiefly to desiccation and high temperatures, it would appear that the ultraviolet spectrum is in itself a factor under certain conditions in the destruction of such ova. LITERATURE CITED Brackett, F. S., and McAuister, E. D. 1932. A spectrophotometric development for biological and photochemical investigations. Smithsonian Misc. Coll., vol. 87, no. 12, pp. I-7, Sept. 26. Brown, Harotp W. 1927. Studies on the rates of development and viability of the eggs of Ascaris lumbricoides and Trichuris trichiura under field conditions. Journ. Parasitol., vol. 14, no. 1, pp. 1-15, Sept. CALDWELL, FRED C., and Errrepa L. 1928. Preliminary report on observations on the development of ova of pig and human Ascaris under natural conditions, and studies of factors influencing development. Journ. Parasitol., vol. 14, no. 4, pp. 254-260, June. CosLENTz, W. W., and Stair, R. 1931. Measurements of extreme ultra-violet solar radiation using a filter method. Bur. Standards Journ. Res., vol. 6, no. 6, pp. 951-976, June. HeErvTet, EF. 1904. Uber die Beeinflussung des Organismus durch Licht, speziell durch die chemisch wirksamen Strahlen. Zeitschr. allg. Physiol., vol. 4, pp. 1-43. MANALANG, C. 1927. Observations on the development of Ascaris ova. Philippine Journ. Sci., vol. 33, no. 3, pp. 249-255, July. Notr, L. O. 1932. Experimental studies on certain factors influencing the development and viability of the ova of the human Trichuris as compared with those of the human Ascaris. Amer. Journ. Hyg., vol. 16, no. I, pp. 288-322, July. Ounsa, T. 1926. On the growth of the eggs of Ascaris lumbricoides. Japanese Journ. Zool., vol. 1, no. 4, pp. 121-122, Mar. 31. Orro, G. F. 1929. A study of the moisture requirements of the eggs of the horse, the dog, human and pig ascarids. Amer. Journ. Hyg., vol. I0, no. 2, Pp. 521-525, Sept. OweEN, WILLIAM B. 1930. Factors that influence the development and survival of the ova of an ascarid roundworm Tovocara caiis (Werner, 1782) Stiles, 1905, under field conditions. Univ. Minnesota Agr. Exp. Sta., Techn. Bull. 71, pp. 1-25, Sept. Ross, RoNALD. : 1916. The life history of Ascaris lumbricoides. (Letter to Editor, dated July 1.) British Med. Journ., vol. 2, no. 2807, pp. 60-61, July 8. NOS I IRRADIATED ASCARID OVA—WRIGHT AND McALISTER 13 Ruppert, W. 1924. Empfindlichkeitsanderungen des Ascariseies auf verschiedenen Stadien der Entwicklung gegeniiber der Einwirkung ultravioletter Strahlen. Zeitschr. wiss. Zool., vol. 123, no. 1, pp. 103-155, June 2. ScHLEIP, W. 1923. Die Wirkung des ultravioletten Lichtes auf die morphologischen Bestanteile des Ascariseies. Arch. Zellforsch., vol. 17, no. 3, pp. 289-366, Aug. Io. ScHwartz, BENJAMIN. 1922. Observations on the life history of Ascaris vitulorum, a parasite of bovines in the Philippine Islands. Preliminary paper. Philippine Journ. Sci., vol. 20, no. 6, pp. 663-669, June. SEIDE, JAKOB. 1925. Zur Kenntnis der biologischen Strahlenwirkung. Untersuchungen am Ascaris-Ei mit ultravioletten, Rontgen-und Radiumstrahlen. Zeit- schr. wiss. Zool., vol. 124, no. 2, pp. 252-304. Stevens, N. M. 1909. The effect of ultra-violet light upon the developing eggs of Ascaris megalocephala, Arch. Entwicklungsmech. der Organismen, vol. 27, no. 4, pp. 622-639, June 15. TSCHACHOTIN, SERGEI. 1912. Die mikroskopische Strahlenstich Methode eine Zelloperations- methode. (Vorlaufige Mitteilung.) Biol. Centralbl., vol. 32, no. 10, pp. 623-630, Oct. 20. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 2 MUD SHRIMPS OF THE ATLANTIC COAST OF NORTH AMERICA (Wir Four PLATEs) BY WALDO L. SCHMITT Curator, Division of Marine Invertebrates, U.S. National Museum (PUBLICATION 3292) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION FEBRUARY 15, 1935 The Lord Galtimore Press BALTIMORE, MD., U. 8S. As MUD SHRIMPS OF THE ATLANTIC COAST OF NORTH AMERICA By WALDO L. SCHMITT Curator, Division of Marine Invertebrates, U. S. National Museum (Witu Four Piates) Except for the comprehensive and invaluable papers of de Man,’ and the exceedingly useful, though less exhaustive, account of Bor- radaile,’ there is for American workers no readily available taxonomic information concerning the several species of Callianassa occurring on the Atlantic coast of North America. To fill this need in part at least, these brief notes and diagnostic key have been compiled. To render them as complete as possible at this time, it has been found necessary to describe three new species, together with a new variety of one of them, and to propose a new name for a specimen that had been assigned to an old, inadequately described species, which to this day has not been certainly rediscovered : Page Galhanassan(Gallichivus) aslagrande ence aes cect oe ase elas aleve ey ifs TOREOTEE TD Ge Oo. BED Toa cle SOC ncn cs POO EE ye) jamaicense vat. lowisianensis............+..+- 4, 12 ANANTH OEM te ey CER ofc. OCC COG ODIO CTD GCOS Als its hartmeyeri, new name for Glypturus grandimanus Balss.............. 4 It is possible also to report a new record of occurrence in Jamaica for Callianassa (Callichirus) longiventris, a species not heretofore seen since the original types were described by A. Milne-Edwards from Martinique. Moreover, in the light of the studies contributory to this paper, Stimpson’s genus Glypturus appears no longer tenable, and its unique species therefore reverts to the genus Callianassa. * A contribution to the knowledge of twenty-two species and three varieties of the genus Callianassa Leach. Capita Zoologica, vol. 2, pt. 6, pp. 1-56, pls. I-12, 1928. The Thalassinidae and Callianassidae collected by the Siboga-Expedition with some remarks on the Laomediidae. Siboga-Exped. Monogr. 39a°, pp. 1-187, pls. I-20, 1928. *On the classification of the Thalassinidea. Ann. Mag. Nat. Hist., ser. 7, vol. 12, pp. 534-551, 1903. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 98, No. 2 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 A word of appreciation is here extended to some of the students and staff of the Biological Laboratory of the Louisiana State University at Grand Isle, La.; in particular, Dr. Ellinor H. Behre, Director, and William W. Anderson, a student of several summers ago, whose in- dustry first brought to my attention specimens of two undescribed species found at Grand Isle, in the determination of which the other information here presented was brought together. I also wish to thank, among others, Drs. E. A. Andrews, of Johns Hopkins University ; C. B. Wilson, of the State Teachers College at Westfield, Mass. ; and A. S. Pearse, of Duke University, Durham, N. C. who have con- tributed specimens of Callianassas to the collections of the United States National Museum. Three west Atlantic species are not included in the key given below: (1.) Callianassa grandimana Gibbes (Proc. 3d. Meet. Amer. Assoc. Adv. Sci., p. 194, 1850. Type locality, Key West). Many attempts have been made to establish this species, regrettably so briefly described by its author. The species to which it seems to stand nearest is C. branneri (Rathbun). Of this latter species I have the greater part of several large specimens taken by Dr. A. S. Pearse on Long Key, Dry Tortugas, Fla., out in the Gulf of Mexico, roughly 60 miles due west of Key West, the type locality of C. grandimana, but until we know more of the Callianassas of our southern States we should refrain from making use of Gibbes’ specific name. Balss some time ago assigned a specimen from Kingston, Jamaica, to C. grandimana, but the very spiny armature of the ventral border of the ischium and merus of the larger cheliped of this specimen definitely precludes any such identity. The ischium of C. grandimana, second segment of Gibbes,’ has “ dis- tant granules on its lower edge,” and the merus, “ the third segment [, | is broader, dilated so as to form below a sharp serrated edge, which is truncated as it approaches the posterior articulation.” Balss’ specimen, moreover, shows a prominently trispinose front ; Gibbes says nothing on this score, but I do believe if the front had been different in any marked degree from that of C. major he would have made some com- ment to that effect ; his observations, so far as they have been checked, have always been proved accurate. His fault was brevity of descrip- *De Man, Capita Zoologica, vol. 2, pt. 6, p. 19, 1928, does say that Gibbes did not describe the merus and ischium, but in the very relations of the several joints of the major cheliped as set forth by Gibbes, the second and third joints are none other than those particular joints; the shape ascribed by him to each of them makes their identification unmistakable. NO. 2 MUD SHRIMPS—SCHMITT 3 tion. Just recently his Squilla neglecta, characterized in remarkably few words, has been found again and recognized without difficulty.’ The front of his grandimana surely cannot be very different from that of C. major or very unlike that of C. branneri. That several authors have assumed that C. grandimana had a trispinose front seems to be due to a mistaken impression based on Stimpson’s passing comparison of the description, not specimens, of Gibbes’ species with his Glypturus acanthochirus. De Man was well aware that Stimpson, as he says, had never seen the Gibbes species, yet he, too, without any good reason, was moved to consider that it might be related to the trispinose C. longiventris. For Balss’ species the name of the collector, Dr. R. Hart- meyer, is proposed: Callianassa hartmeyeri. (2.) Callianassa siguanensis (Boone). (Glypturus siguanensis, Bull. Bingham Oceanogr. Coll., vol. 1, art. 2, p. 85, fig. 17, 1927. Type locality, Siguanea Bay, Isle of Pines.) This species would key out below with C. branneri (p. 4, below). For want of evidence other than that given by the author I am inclined to consider it identical with this last-named species. (3.) Callianassa occidentalis Bate (Rept. Challenger Macrura, p. 29, pl. 2, fig. 2k, 1888. Type locality, off Sombrero Island). Only the larger left cheliped is known, so the species ( ?) cannot be satisfactorily keyed out ; moreover, it may even be the cheliped of a larger specimen of the “ Cheramus ” occidentalis of the same author noted in the key below (II. A., p. 5), a considerably mutilated specimen lacking the chelipeds. KEY TO SPECIES OF CALLIANASSA I. Telson broader than long, third maxillipeds more or less broad and flattened, especially ischium and merus, and often the propodus too. A. Lateral angles or projections of front not spined. 1. Inner uropods narrow, about four times as long as broad, styliform or straplike; rostral projection low, blunt, or rounded triangular. a. Carpus of larger cheliped more than four times as long as its great- est width; merus with a shallow projection or low granulated tooth on ventral border; ischium with a long, prominent hooked tooth or spine near middle of ventral border... .islagrandce, n. sp. b. Carpus less than three times as long as wide; merus with strong tooth near proximal end of lower border; ischium not known (North Carolia toniiorida)c2 ssc ceists + 6 ses'ei ss ele « major Say.” *“Lunz, G. Robert, Jr., Charleston [S. C.] Museum, Leaflet No. 5, pp. 1-8, 1933. *Journ. Acad. Nat. Sci. Philadelphia, vol. 1, pt. 2, p. 238, 1818; de Man, Capita Zoologica, vol. 2, pt. 6, p. 30; pl. 7, fig. 14-14b; pl. 8, fig. 14c-14d, 1928. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 2. Inner uropods wider, either (a) about twice as long as wide or (b) not much longer than wide. a. Inner uropods about twice as long as greatest width; rostrum a conical acuminate spine. i. Merus of larger cheliped armed with a toothlike process near proximal end of ventral border; upper border of carpus about as long or longer than upper border of palm. . jamaicense, n. sp. ii. Merus unarmed below; upper border of carpus not exceeding three-fourths the length of upper palmar border (Puerto Rico and:Batbados)s aces sscmuctansm one ae marginata Rathbun.° b. Inner uropods not much longer than wide; rostrum low, triangu- lar, subacute, or rounded off. i. Merus of larger cheliped armed with a strong tooth near proxi- mal end of lower border; upper border of carpus three-fourths or more the length of the upper palmar border (Massachusetts tom Nonthe Carolina) aseriscereeeee cies atlantica Rathbun." ii. Merus inconspicuously serrate or denticulate, but without strong tooth below; carpus from a half to two-thirds as long as upper border of palm (Brazil, Bermuda, Puerto Rico, Barbados, and the Dry, dortugas, Plorida)). esses branneri Rathbun.* B. Frontal margin trispinose. 1. Both chelae armed with three strong spines above; upper border of carpus three-fourths the length of the palmar border; carpus strongly spined on lower border, merus spined above and below (Hloridasamaicas Barbados) meee acanthochirus Stimpson.* 2. Chelae not armed with strong spines above. a. Eyestalks flattened, cornea on dorsal surface ; merus of larger cheli- ped multidenticulate below, of minor cheliped unarmed; upper border of carpus of larger cheliped less than half the length of thesupper palmar bordeGeesane see cece tes rathbunae, n. sp. b. Eyestalks subcylindrical, cornea terminal or nearly so. i. Merus of large cheliped with more than five spines on ventral border, of minor cheliped unarmed; upper border of major carpus less than half as long as upper palmar border (Mar- tiniquesand Jamaica) aemeceeteeeee longiventris A. M.-Edw.” ii. Merus of large cheliped with four or less spines below, of minor cheliped with one or two; upper border of major carpus more than half the length of the upper palmar border (Jamaica). hartmeyeri, new name.” ® Bull. U. S. Fish Comm. for 1900, p. 92, fig. 15a-d, I90T. 7™U. S. Nat. Mus. Bull. 138, p. 107, footnote, 1926; C. sttmpsont Smith, Rep. U. S. Fish Comm. for 187, p. 549, pl. 2, fig. 8, 1873; de Man, Capita Zoologica, vol. 2, pt. 6, p. 37, pl. 9, figs. 17, 17d, 1928. § Proc. Washington Acad. Sci., vol. 2, p. 150, pl. 8, figs. 5-8, 1900. > Glypturus a., Ann. Lyceum Nat. Hist. New York, vol. 10, nos. 4, 5, p. 121, 1871. ” Nouv. Arch. Mus. Mem., vol. 6, p. 92, 1870. De Man, Capita Zoologica, vol. 2, pt. 6, p. 24, pl. 6, figs. 12, 12h, 1928. 4 Glypturus grandimanus Balss, Zool. Anz., vol. 61, p. 179, 1924. NO. 2 MUD SHRIMPS—SCH MITT 5 II. Telson longer than broad; third maxillipeds narrow; frontal margin without lateral spines; eyestalks flattened, cornea dorsal or lateral; inner uropods less than twice as long as wide. A. Telson with a pair of spines on the posterior half of either lateral margin, and a spine at middle of posterior margin; rostrum a little shorter than the eyestalks; chelipeds and external maxillipeds unknown (off Som- (ARES VIG) cially Lier ae Se ear A een a a batei Borradaile.” B. Telson laterally with several spinules, posteriorly unarmed; upper border of carpus less than half as long as upper palmar border ; rostrum longer than eyestalks; external maxillipeds narrow (Puerto Rico). minima Rathbun.” CALLIANASSA (CALLICHIRUS) ISLAGRANDE, n. sp. A distinctive species. Lineae thalassinicae distinct ; cervical groove crosses the carapace at nearly one-fourth the length of the carapace from the hinder margin. Rostral projection of front low triangular, extending forward beyond the level of the blunt, inconspicuous lateral projections by about one-fourth or one-fifth its width measured be- tween those projections; the rostrum reaches forward less than one- third the length of the eyestalks, and these in turn reach between one- fourth to one-third the length of the second segment of the antennular peduncle. The eyestalks are dark brown, with black corneae, and are about twice as long as broad ; their terminal one-third to two-fifths are each drawn out into a slender spine slightly curved upward, and with the tip very slightly exerted ; inner borders of eyestalks contiguous to about the level of the anterior margins of the corneae ; the latter are opposite the distal end of the second segment of the antennal peduncle. Fourth segment of the antennal and second segment of the antennular peduncles reach about equally far forward; fifth or terminal segment of antennal peduncle reaches about two-fifths the length of the distal segment of the antennular peduncle. Right cheliped of male much the larger; remarkably long and slender, quite the most striking character of this species; the carpus exceeds a little the propodus plus the dactyl, even when the latter is extended straight forward ; carpus and palm about equally wide ; carpus nearly four and one-half times as long as its greatest width; palm shorter, measured on the upper margin, about three and one-half times ® Ann. Mag. Hist., ser. 7, vol. 12, p. 546, 1903. Cheramus occidentalis Bate, Rept. Challenger Macrura, p. 32, pl. 2, fig. 1, 1888. Callianassa batei de Man, Capita Zoologica, vol. 2, pt. 6, p. 10, pl. 1, fig. 3, 1928. 18 Bull. U. S. Fish Comm. for 1900, p. 92, fig. 16a-d, 190T. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 as long as wide; merus about four-sevenths the length of carpus; ischium a little longer than merus and about two-thirds the length of carpus. Ischium granulated with low, rounded, “ pearly ” granules on both faces, more sparsely on outer than inner faces, forming a denticulated inferior margin; upper margin evenly concave, longitudinally deeply grooved; upper part of outer face of joint, including outer ridge, forming dorsal groove without granules, smooth and shining; inner face of ischium more heavily granulated than outer; just before the middle of its length there is a long, prominent, well-granulated tooth on the lower margin of the ischium, a tooth about as long as the ischium is wide. Upper margin of merus unevenly and shallowly concave in lateral view ; at about the posterior third of its ventral margin is a low granu- lated tooth, appearing to be made up of sharper, more prominent and more crowded granules than those ornamenting the sides of the ischium ; upper border longitudinally grooved like ischium ; outer ridge forming groove also very smooth. The carpus is approximately parallel-margined, though finding its greatest width at about the distal end of the proximal three-sevenths ; it is smooth and shining on inner and outer faces but finely denticulate on the upper margin for the greater part of its length, the denticles becoming obsolescent close to its posterior extremity ; lower margin of carpus keeled, keel less conspicuous and more or less replaced by rather widely separated denticles or granules in anterior four-sevenths and also obscurely so at the hinder angle. Palm, like carpus, smooth and shining on lateral faces, upper margin on inside very finely denticulate, lower keeled, keel smooth; on either side of keel there are tufts of long hairs, toward lower edge of palm the hair tufts are roughly alternate on either side of this keel; along the outer side of the keel from proximal end to tip of fixed finger there may be counted about eight tufts, on inner side eleven, the last five of which are more on the “ ventral’ surface of the finger, for here the keel seems to become less distinct anteriorly and the finger is a bit flattened below on the inner side of the palm; the keel of the lower margin is paralleled by a row of denticles for a little more than its distal half, denticles disappearing on the inner, lower margin of the finger where a blunt keel takes their place; the last hair tuft just below the extreme tip of the movable finger is elongate and more bushy than the other tufts. The movable finger for the greater part is gently bowed, but at the far end is abruptly hooked; below, it is armed proximally with a ———————— NO. 2 MUD SHRIMPS—SCH MITT 7. prominent blunt tooth, and anterior to it, with two blunt denticles ; distally on the outer, upper border of the terminal hook of the movable finger there is a conspicuous blunt right-angled tooth topped with a thick tuft of hair. This upper, outer tooth on the movable finger makes the digit appear terminally bifurcate ; the tooth seems to be a feature characteristic of this species alone ; on the upper margin near the base of the tooth is a blunt tuberculiform tooth, and beginning a little distance behind this on the inner side of the upper margin of the finger, a row of low tubercles, about seven in the type; they seem not to be developed in smaller specimens ; the fixed finger arises some distance— about half its length—behind the distal margin of the palm and forms with the forward projection of the palm a deep sinus; the fingers, when closed, have a large gape between them, half of which is formed by the sinus just described ; the terminal portion of the movable finger closes or hooks outside the tip of the fixed finger. The smaller cheliped, as compared to the larger, is slight and thin, carpus and propodus together being noticeably shorter than the carpus alone of the larger cheliped ; the ischium is fairly straight, unarmed, thin, and flat, a little longer than the merus, shorter than the carpus, and a little shorter than the chela to the tips of the fingers when closed ; from its hinder border the carpus narrows anteriorly in lateral aspect, likewise the hand to the tips of the fingers ; measured to the tip of the extended movable finger, the hand and carpus are subequal ; measured at the middle of its length, the width of the carpus is contained twice in the length of its upper border; the palm, measured from the base of the sinus between the fingers, is longer than wide and longer than the fixed finger measured from the same point, but equal to the movable finger; all joints of the cheliped are smooth and shining ; merus and carpus hairy on lower margins, palm thickly so on both margins, fingers on outer surface face adjacent to prehensile edges provided with a thick felt or pad of hairs; prehensile edge of either finger denticulated, armed with about 15 small, sharp, triangular teeth, of which the most anterior are rather small; distal fourth of either prehensile edge or margin more or less without teeth. The inner face of the ischium of the third maxilliped (pl. 3, fig. 2) shows a short crescentic row of tiny granulations, scarcely to be ob- served with a glass in a wet specimen; continuing the curve of the crescent formed by not over a dozen of these granulations is a slight ridge, set off by no more than an incised line beside it, which extends about three-fourths the length of the joint. The abdomen is soft, broad, and depressed ; the first two somites are dorsally thin and membranous, the others are thicker and tougher, 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 symmetrically grooved, and with the exception of the sixth, are orna- mented on the epimeral region of either side with a pad or felt of hair. The telson is about one and one-half times as wide as long, and a little less than half as long as the sixth abdominal somite ; the telson and the sixth somite are ornamented with hair tufts about as in C. major, but otherwise the two are of very different shape. The telson of C. isla- grande is truncate or more or less squared-off posteriorly, and but little contracted anteriorly; medially there is a longitudinal, smoothly rounded, raised area, but lower than the inflated lateral portions of the dorsum of the telson, which is bounded laterally by bowed-out, smooth grooves or depressions confluent behind but separating again to pass either side of a rounded, raised area, the pushed-up median portion of the hind border of the telson; the outer uropod is more or less triangular with but gently convex posterior margin, the inner uro- pod is narrow, tongue- or strap-shaped, distally rounded, and nearly four times as long as wide. In the useful key given by de Man™ our species would take place alongside of C. major, having, like it, very narrow inner uropods, about four times as long as broad, and, to use his expression, almost styliform. However, the different eyestalks, the remarkably elongate larger chela of the male, and the character of the telson sharply differ- entiate C. islagrande from that species. The females need no separate description ; their chelipeds are both much like the minor cheliped of the males. Type.—vU.S.N.M. no. 69362, the largest of 10 specimens (401, 62) collected by William W. Anderson at Grand Isle, La., in the summer of 1930. It is a male, measuring approximately (in millimeters) : carapace 19 long, abdomen 62 exclusive of telson, telson 6 long by 8.5 wide. The larger right cheliped measures: ischium 21, merus 17, carpus 30 (greatest width 6.5), hand on upper margin 19. The ex- treme forward reach of the movable finger from the articulation to the summit of the tooth on the upper outer margin of the finger measures 10; from articulation to tip this finger measures only 8.5. The smaller, left cheliped measures: ischium 9; merus 8; carpus 11, width at middle 4.8; palm, upper margin 5, width at middle 3.5, lower margin to tip of fixed finger 7.3; movable finger from articulation to tip 6. I have also seen a lot of 10 females collected by Dr. Ellinor H. Behre at the same place July 5-17, 1928, and a male taken July 15 of the same year by Mr. Anderson. * Siboga-Exped. Monogr. 39a’, p. iii, 1928. NO. 2 MUD SHRIMPS—SCH MITT 9 CALLIANASSA JAMAICENSE, n. sp. Lineae thalassinicae present; central dorsal “ oval’’ area bounded posteriorly by the cervical groove, and delimited before by a trans- verse groove behind the rostrum; cervical groove crosses the carapace two-sevenths of the length of the carapace from its hinder margin ; the groove crossing the carapace behind the rostrum is about one- seventh the length of the carapace behind the tip of the rostrum; the rostrum extends beyond a line joining the barely marked, obsolescent lateral angles of the front by about half this distance. The rostrum is a stout conical spine reaching about as far forward as the distal margin of the cornea. The eyestalks are contiguous throughout the extent of their inner margins ; on the inner margin just in advance of the centrally placed cornea there is the suggestion of a tubercle reminiscent of the one described by Holmes” for his Lepidophthalmus, now Callianassa (Callichirus) eiseni from Lower California, a species to which the present one shows kinship, yet from which it may readily be dis- tinguished by the proportions and armature of the major cheliped. The eyestalks of C. jamaicense fall a little short of the first segment of the antennular peduncle; the second segment of the antennular peduncle is very little longer than the first measured from the orbital margin of the carapace, and a little short of a third of the length of the third segment; the terminal segment of the antennal peduncle reaches the middle of the last segment of the antennular peduncle, the fourth segment a little past the second of the antennular peduncle, and the second antennal segment and the first or basal antennular segment reach about equally far forward ; the antennal scale is represented by a small, inconspicuous spine at the antero-lateral angle of the second segment ; the first or basal segment reaches about to the middle of the cornea ; as in all these Callianassas, the third segment of the antennal peduncle in dorsal view scarcely appears to be more than an articulation between the second and fourth segments of the normally five-seg- mented peduncle; the third segment is for the most part behind and hidden by the second segment. The ischium of the larger right cheliped is nearly equal to the upper border of the palm in length and a little (about one-fifth) shorter than the merus ; upper border of merus a little longer than that of carpus; merus about one and three-fourths times as long as its median width ; carpus at middle of its length about as wide as length of upper border ; = roe, California Acad! Sci, ser. 3; Zool., vol. 3, no. 12; p. 311; pl. 935; figs: 6-13, 1904. Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 median width of palm and carpus equal; fixed finger measured from the sinus at its base is three-fourths the length of the palm measured back from the same point ; it is fairly long and ends in a stout conical subacute tip, and a very low triangular tooth is situated about the middle of its length. The movable finger is nearly a third longer than the fixed one and armed with three stout conical teeth, a blunt one near the base tending to fit into a notch at the upper end of the base of the sinus between the fingers ; the second and larger of the other two more pointed teeth is placed just before the middle of the finger, and the third and smallest is located about one-third the distance from the second tooth to the tip ; the upper border of the movable finger is smoothly rounded off, but toward the articulation is somewhat eroded-looking, two short grooves and several pits appearing here; there is a low, conical, inconspicuous tubercle, and perhaps a second lower one adjacent to it, just before the articulation ; the fingers cross in such a way that the tip of the fixed one closes between the anterior of the three teeth of the movable finger and its hooked tip, and the two conical teeth of the upper finger bite just before the triangular tooth of the lower margin. On the outer face of the palm, more or less continuing the line of the outer edge of the prehensile margin of the fixed finger back on to the palm, is a well-marked, curved carina extending back nearly half the length of the palm ; above this carina is a low, depressed, triangular area; the anterior margin of the palm is armed with two blunt, for- wardly directed teeth, one forming the outer side of the notch at the upper end of the base of the sinus between the fingers and the second placed laterally just below the articulation of the movable finger. Lower border of the palm and movable finger somewhat cristate and obscurely and bluntly serrate, rather, one might say, undulate, owing to the insertion of hair tufts found there ; above the palm is more or less carinate in its posterior two-thirds. The carpus is cristate above and below, the anterior dorsal and ventral angles forming each a com- pressed triangular tooth. All joints are smooth and shining. The merus shows a blunt longi- tudinal ridge on the outer face; the infero-proximal angle of this joint is armed with a conspicuous, angled lobe or toothlike process ending primarily in a stout, elongate, conical spine, on the inner hinder side of which is a secondary, smaller, and somewhat curved spine; the laminate lower margin of this joint 1s seven-toothed in its anterior two-thirds, the first two flattened, the second of these obscurely doubled, the following five placed on the anterior curve of the lower margin more tuberculiform, though compressed; the upper margin NO. 2 MUD SHRIMPS—-SCH MITT MIE of the merus is interrupted close to the posterior end by a conspicuous narrow U-shaped notch, at the upper extremities of which the dorsal margin of the joint forms a blunt tubercle. The ventral margin of the ischium is, beginning near the hinder end, finely and for the greater part rather evenly denticulate, the denticles, however, gradually increasing in size anteriorly, the last two or three being larger, the last of these very large comparatively, and forming rather a tuberculiform tooth almost half a millimeter long and distally obscurely bilobed ; in advance of this tooth the lower margin is blunt- ridged; above on the outer face, and shorter than the denticulated portion of the lower margin of the ischium, is an obscure, blunt ridge. When both chelipeds are moderately extended to about the same degree, the fingers of the smaller reach scarcely half their length beyond the distal margin of the carpus of the larger. The ischium is about as long as the merus and a little longer than the carpus or upper margin of palm, but only four-sevenths the length of the movable finger. The upper margin of the dactylus is ridged and grooved, the prehensile edges of both fingers are wide and somewhat hollowed out and the movable one is furnished with two anteriorly converging lines of ob- scure, small tubercles; the extreme tips are somewhat abruptly con- stricted to form conical spinous tips so that there is apparently a blunt tooth at the anterior extremity of the outer moiety of each fairly broad prehensile edge ; when closed, the tips of the finger reach about equally far forward; there is a sizable gape between them. The palm is rounded above, cristate below, except on the distal half of the fixed finger, carpus blunt above, cristate below, superior and inferior distal angles subacute, merus blunt-ridged above and below; ischium blunt above, with finely denticulate margin below; the ischium and merus are about of the same length measured on the upper border, either a little longer than the carpus, about one-third longer than the upper border of the palm, and a little less than two-thirds the length of the movable finger. The inner face of the ischium of the third maxilliped (pl. 2, fig. 8) carries a smooth, blunt, yet well-formed, carina wholly without spinous armature, or tiny scalelike excrescences. Telson somewhat rectangular, about one and one-third times as long as wide, little less than half as long as the sixth somite. Posterior margin three-lobed, shallow, median lobe occupying more than one- third of the posterior margin, and the gently curved lateral angles carried evenly and smoothly around to their respective sides to merge with lateral margins of the telson; dorsal surface evenly convex in both directions ; across the dorsum of the telson a little way before the I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 base is a fine, sharply impressed line occupying more than half the total width of the telson. Inner uropod narrow, a shade more than twice as long as its greatest width measured on, and at right angles to, the median axis drawn from articulation to apex of the blade. Basal joint of uropod armed with a sharp, stout spine; there is a similar but even longer spine near the proximal end of the inner border of the outer uropod, forming with the blade a notch into which the outer margin of the inner blade snugly fits. Holotype-—U.S.N.M. no. 69363, male, taken from a brackish pond at Montego Bay, Jamaica, by C. B. Wilson, June 29, 1910. It measures (in millimeters) approximately 73 long, tip of rostrum to end of telson ; somewhat broken carapace about 18+, abdomen and telson 55, telson 6 long by 7.5 wide. Large chela, lower border of palm to tip of fixed finger 21, upper border 13.5, dactylus 12.5, width of palm at middle of length 12.5, upper border of carpus 12.5, width 12.5, upper border of merus 13.7, ischium 11. Small chela, lower border of palm to tip of fixed finger 16, upper border 6.1, dactylus 12.5, width 8.3, carpus 7.2 long, width at middle of length 6.3, length of merus 8, ischium 8. A smaller specimen of about 61 mm in length, also from Montego Bay, taken on June 24 of the same year by Dr. E. A. Andrews, likewise from a salt-water pond, is like the type in all particulars, except in the ventral armature of the merus of the large cheliped, which is five- instead of seven-toothed toward the distal end of the joint. The fine, sharply impressed line across the midportion of the base of the telson is present, but does not extend as far to either side as in the type. CALLIANASSA JAMAICENSE var. LOUISIANENSIS, n. var. From Grand Isle, La., comes another specimen very similar to the preceding, which I should almost have been tempted to describe as a distinct species but for the fact that its larger right hand shows some evidence, though slight, of having been injured. For the present at least it had best be considered as no more than a varietal form. There is only a single male specimen taken by Chenier Ronaville, July 18. 1928. Type—vU.S.N.M. no. 69364. The description of the type of the species proper as regards carapace, rostrum, and frontal appendages, about fits the variety ; the eyestalks, however, have a very pronounced tubercle on the inner margin, whereas only a suggestion of one was to be seen in the type of the species. It was only mentioned in con- nection with the type, because a very definite though small tubercle is NO. 2 MUD SHRIMPS—SCH MITT 1033 visible in the only other specimen of the typical species at hand, also from Montego Bay, Jamaica. The major right cheliped does not differ much from that in the typical species in the proportions of its joints and their armature ex- cept in the fingers ; merus and carpus about subequal, measured on their upper borders; merus about twice as long as its median width; fixed finger from hind end of sinus between the fingers is one-half the length of the palm measured back from the same point. The fixed finger is short and strong and ends in a stout, conical, up-turned ex- tremity and has a strong, blunt, tuberculiform tooth situated at about one-third the length of the finger from the back of the sinus between the fingers to the tip. The movable finger had the tip broken off before capture and may have lost from a tenth to a fifth of its length, the part remaining slightly exceeding in length the fixed finger. On the cutting edge it is provided with two teeth, a stout conical one near the base of the finger tending to fit into a notch at the upper end of the base of the sinus between the fingers, and an anterior longitudinally elon- gated laminate tooth; these teeth are so spaced that the tooth on the fixed finger bites between them, while the tip of the fixed finger closes in advance of the laminate tooth of the movable finger ; on the upper margin of the movable finger is a single blunt tubercle, more con- spicuous than either of the tubercles similarly placed in the type of the species ; also in the variety the finger is smooth above and not eroded toward the articulation; in the second of the typical specimens the finger, again, is eroded and the tubercle inconspicuous. Near the hinder end of the carina forming the posterior two-thirds of the upper border of the palm there is a brief emargination, followed by a small tooth; that this effect may have been brought about by an injury is possible ; the conspicuous carina on the outer face of the palm of the type of the species is here represented by an inconspicuous impressed line perhaps two-sevenths the length of the palm in line behind it ; no depressed area is evident above this line. The antero-inferior angle of the carpus shows two tiny tuberculi- form projections, one either side of a hair tuft at that angle. The ventral margin of the merus, other than the two-spined lobe at the posterior end, as in the type of the species, is unarmed except for three compressed teeth near the anterior end of the middle third of the margin; the merus of the smaller of the two typical specimens closely resembles that of the varietal form, though it has five teeth in place of the three just mentioned. Posteriorly, the upper margin of the merus has a shallow emargination in place of the very definite notch found in the typical specimens. The tooth at the anterior end of 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 the denticulations arming the ventral margin of the ischium is larger than that in the type of the typical species, being nearly half a milli- meter long. The smaller cheliped is not much different from that of the type of the species proper, the fingers are more slender and the hand more triangular ; the tip of the movable finger is also wanting. The smooth, blunt ridge on the inner face of the ischium of the third maxilliped (pl. 2, fig. 7) appears unarmed or unadorned ; only by careful inspection, and then with a glass, can five little scales or tiny, scarcely perceptible, well-separated, flattened spines be observed in line on the proximal third of the ridge. Next to the differences from the typical specimens in the large chela and fingers, those found in the telson and the uropods are the most marked. The telson is even more rectangular-appearing than that of the type of the species, as the lateral angles are not so broadly rounded and the hinder margin is more truncate or straight ; it is, however, like that type in being about one and one-third times as wide as long, and contained in the length of the sixth abdominal somite about twice ; the telson is transversely evenly, though shallowly, convex, flatter than in the typical species, but unlike it, has the median longitudinal con- vexity interrupted by a not inconspicuous depression just before the hinder margin ; the inner uropod measured as in the type of the species proper is but one and one-half times as long as wide ; the basal segment of the uropods is armed with a low, blunt, triangular spine, and in place of the conspicuous spine on the proximal margin of the outer blade of the typical specimens there is an inconspicuous, low tubercle ; no fine impressed line appears across the middorsal region of the telson parallel to and a little before the basal margin ; there is a short, not at all well-marked, shallow, transverse depression on either side of the midline, but not crossing it; these two inconspicuous depressions are well separated from one another transversely and are in no way like the very definite fine line of either of the typical specimens, which very distinctly crosses the middorsal line of the telson. The differences are such as one might expect, in part at least, be- tween male and female specimens, but I can find no evidence that any of the specimens are female. True enough, the type retains but one basally much mutilated third leg, but there are in the other two speci- mens no traces of external female apertures, and the first pleopods of all three are of the same masculine character. This varietal specimen measures (in millimeters) approximately 68 long, carapace and rostrum 16, abdomen and telson about 52; telson alone about 4.9 long by a shade over 6 wide. The large chela, measured —— - NO. 2 MUD SHRIMPS—-SCHMITT T5 on lower border to tip of fixed finger, is 15.5 long; upper border of palm to articulation of dactylus 10, broken dactylus 7.7, width of palm at middle of length 11, length of upper border of carpus, as well as width, 11, upper border of merus 10.3, ischium 8.6. Small chela, movable finger 10, upper border of palm 4.3, width 6.5, lower border to tip of fixed finger 12, carpus 5.7, width at middle of its length the same, merus 6, ischium 6.5. Remarks—In giving a name to this proposed new variety, I am reminded of the varietal form that de Man” has indicated for Borra- daile’s C. (Callichirus) novae britanniae, and again of his discussion ™ ‘of C. (Tryphaea) californiensis var. japonica Bouvier and C. (T.) japonica (Ortmann) and his C. (Callichirus) longiventris var. borra- dailei.* I believe the specimen worthy of varietal designation until such time as additional material may call for a change of opinion. CALLIANASSA (CALLICHIRUS) RATHBUNAE, n. sp. With tridentate front and the “ oval ” area before the cervical groove scarcely delimited from the front, although a very faintly indicated groove crosses the anterior part of the carapace at a distance behind the orbital margin about equal to the length of the rostrum; laterally, this faint groove trends backward and downward toward the linea thalassinica on either side but does not join either it or the cervical groove behind. The cervical groove is deep and crosses the dorsum of the carapace about two-sevenths of the length of the carapace in advance of its hinder margin. The rostrum is a slender short spine reaching about one-half the length of the contiguous portions of the inner margin of the eyestalk. The latter are in contact for about two- thirds of their length, flaring in the anterior third widely outward to terminate each in a little pointed tubercle at the juncture of the inner border of the eyestalk and the gently incurved outer border in front of the cornea. This tubercle lies in line with the midpoint or center of the cornea and about the length of the corneal diameter in front of it. The corneae are light brown and occupy the outer two-fifths of the width of the stalk and approximately the fourth and fifth sevenths of the length of the stalk. The eyestalks just overlap the distal margin of the basal segments of the antennular peduncle; thus the first segment of this peduncle is not visible except for a small portion either side of the eyestalks ; * Capita Zoologica, vol. 2, pt. 6, p. 49, 1928. ™ Idem, pp. 18, 10. “dent, p. 27. 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 the second segment of the peduncle is about four-sevenths the length of the first, and in turn three-fifths the length of the terminal segment. The second segment of the antennal peduncle on the right side reaches about as far forward as the basal segment of the antennular peduncle ; the fourth segment reaches nearly to the distal margin of the terminal segment of the antennular peduncle; the terminal segments of both peduncles are nearly subequal in length and each about two-thirds the length of the fourth segment of the antennal peduncle; the antennal peduncle of the left side seems to be placed lower, as the distal margins of its several segments each fall short in turn of the forward distance attained by the segments of the extended peduncle on the right side. The lateral frontal spines are slender, sharp, incurved, and to some degree movable, as they are joined to the front by a somewhat flexible noncalcified extension of the carapace ; they lie just without the line of the inner margin of the antennal peduncles. The lineae thalassinicae diverge more widely anteriorly than ‘posteriorly, and so at the front the “oval” area which they appear laterally to limit seems much wider there than posteriorly, thus giving the dorsum of the carapace of this species an inverted wedge-shaped look unlike the other species described in this paper. The large left cheliped has a very short carpus, which is a little short of being twice as high as long. Measured on the upper border, the palm is very slightly longer than the merus, and a very little shorter than the movable finger, about twice the length of the carpus and about one-fifth longer than the ischium; the palm is about as wide at its midpoint as the upper border is long. The ischium is armed beneath the anterior end with three moderately long, slender spines, toward the posterior end also with three more widely spaced, much smaller, slender ones, and at a point midway between the two groups there is a single spine of intermediate length; all of the spines are directed forward, as are those arming the lower border of the merus; on the latter are to be counted 12, which get smaller anteriorly, with the exception of the second spine, close by the first one, which seems smaller than it should be, perhaps abnormally so. The carpus and chela are smooth and shining; the upper and lower margins of the former are thin, laminate, and turned over to form a serrate edge above and below ; in the ‘case of the carpus for the greater part of the hinder margin this serrate edge is so turned over as not to be visible in the outer, lateral view of the cheliped, except for the most anterior two or three serrations at the infero-distal angle; the upper and lower margins of the palm are cristate and also serrate, but the serrations are not so much turned at right angles to the perpendicular as in the NO. 2 MUD SHRIMPS—SCH MITT WZ carpus, for, although the dorsal serrations are scarcely visible from the side, the ventral ones are plainly so; the latter run forward only about half the length of the ventral border of the fixed finger. The prehensile edge of the dactylus has two elongate, not very prominent, laminate teeth, the fixed finger a single low triangular, more or less inconspicuous one a little behind the middle of its length. In the smaller chela the ischium is very little longer than the merus, subequal ‘to the carpus, and nearly a third longer than the upper border of the palm ; the movable finger is a little longer than the palm above. The greatest width of the carpus is contained about twice in its length, and equal in width to the palm; the width of the latter is contained in its upper border about one and one-third times. On the inner surface of the ischium of the third maxilliped (pl. 3, fig. 1) there is a curved row of 17 teeth, of which the proximal 7 are strong, acute spines, and the remainder on the distal moiety of the comb are dentiform rather than spiniform. Telson truncate, hinder margin slightly scalloped, a little less than one-half the length of the sixth somite and about four-fifths as long as its greatest width ; inner uropod longer than the telson, but shorter than the outer uropod, broadly triangular; outer blades but weakly sculptured. Somites of the abdomen remarkably smooth and without ornamentation. Type.——vU.S.N.M. no. 23010, the larger of two male specimens in the collections of the National Museum, both from Bluefields, Jamaica, where they had been washed up after a storm in the summer of 1899. It measures (in millimeters) : carapace and rostrum 18 long, abdomen and telson together 43, telson 5.3 by 6.7 wide; large cheliped, ischium to long, merus I1, carpus 6.5 long, greatest width 10.5, palm, upper border 12 long, movable finger 12, lower border 22.2, width of palm at middle of length 11. Small chela has the ischium 8.3 long, merus 7.7, carpus 9 by 4 wide, palm, upper border 6, movable finger 6.8, lower border, including fixed finger, 12.0, width of palm at middle of length 4.3 CALLIANASSA (CALLICHIRUS) LONGIVENTRIS A. M.-Edw. Callianassa longiventris A. Milne-Edwards, Nouv. Arch. Mus. Mem., vol. 6, p. 92, 1870. Callianassa (Callichirus) longiventris de Man, Capita Zoologica, vol. 2, pt. 6, p. 24, figs. 12, 12 h, 1928. From the bathing beach at Montego Bay, Jamaica, Dr. E. A. Andrews, of Johns Hopkins University, took two specimens of a 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Callianassa, one of which regrettably lacks the larger chelipeds, for both represent an exceedingly rare species known heretofore only from the type lot of four described in 1870 by A. Milne-Edwards, from Martinique. A comparison with the detailed description given by de Man for one of the specimens of the type lot reveals certain small differences, mention of which might not be out of place here. The rostrum is longer, extending about as far forward as the tuber- cles tipping the eyestalks; the acuminate, lateral spines of the front reach about as far forward as the corneae and so are longer, too, than those of the type. I should say that the eyestalks are subcylindrical, though they appear in dorsal view to be more or less cylindrical ; the corneae are rounded, bulged-up prominences on the anterior dorsal surface of the trans- versely convex stalks; the patches of dark pigment are concentrated within in the median, posterior halves of the corneae. The first and third segment of the antennular peduncle are about subequal, each a little longer than the second segment. The terminal segment of the antennal peduncle exceeds the corresponding segment of the antennular peduncle by about three-fourths its length ; the fourth or penultimate segment reaches about two-thirds the length of the second segment of the antennular peduncle; while the distal dorsal margin of the third, which is about all that is visible of it from above, about attains the distal margin of the basal segment of the antennular peduncle. The telson is rather angulated, posterior margin rather straight with the triangular median prominence well marked; medially, in the an- terior half above, there is a small, smooth and shining, more or less semicircular raised area from which the telson falls away to its mar- gins ; two obscure ridges run back to the postero-lateral angles of the telson ; on the posterior margin just inside either of these angles is a tuft of long hair; the hinder margin of the raised area on the dorsum of the telson is also furnished with a pair of these hair tufts. The larger cheliped, as in the type described by de Man, is also on the left side; of the ventral ischial spines, the last is the larger, and so it seems to be also in the drawing that de Man gives of this joint; on the lower border of the merus there are but three instead of four spines, the first the smallest, the third larger and more forwardly directed than the others; these three, moreover, are all placed close EE NO. 2 MUD SHRIMPS—SCH MITT 19 together at the infero-posterior angle of the joint and not so placed as to occupy the greater portion of the ventral margin as shown in the picture of the variety figured by Borradaile” and referred to by de Man. Measurements (in millimeters) are as follows: carpus on the upper border 6 long and 8 broad at the distal margin; palm 11.5 on the upper margin, 9 broad at the middle, and 17.5 on the ventral margin to the tip of the fixed finger ; movable finger from articulation to tip 9 long. The distal border of the palm is distinctly crenulate and, as in the type, is separated by a minute triangular incision from a low triangular tooth which, in the case of our specimen, has its apex at about a fourth of the length of the cutting edge of the finger from the small triangular notch at the base to the tip; the cutting edges of both fingers are sharply cristate throughout, biting so as to about impinge on the “ crown”’ of the tooth of the fixed finger; there is a low, not very conspicuous, triangular tooth on the movable finger. In the smaller cheliped, the ischium is armed below with Io instead of 7 spines ; the merus has a spine as in the type; it is here placed on the inferior margin at the end of the first third, but, unlike the type, shows a second spine midway between it and the infero-posterior angle of the merus; of this spine the distal half has been broken off, apparently before capture. Measurements (in millimeters) are as follows: carpus 6 long and 5.7 wide at the distal end; upper border of the palm 7 long, the lower to the tip of the fixed finger 14, width at middle 6; the distal margin of the palm is crenulate, but not so distinctly so as in the larger chela, the fingers are armed much as are those of the larger hand; the movable finger is about 8.8 long. The comb on the inner surface of the ischium of the third maxilliped (pl. 3, fig. 3) is armed, in the distal half with 12 to 15 small sharp spinuliform teeth, followed in its proximal portion by three large, strong, rather widely separated spines. Of this more complete specimen, a male, the carapace and rostrum together are about 20.7 long, the abdomen and telson together ap- proximately 63; telson is about 6.2 wide at base by 5.8 long. The specimen without chelipeds is a female, a little larger in size, measuring not less than 88 from tip of rostrum to end of the telson. ” The fauna and geography of the Maldive and Laccadive Archipelagoes, vol. 2, pt. 3, p. 752, pl. 58, fig. 2, 2b, 1904. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 CALLIANASSA (CALLICHIRUS) ACANTHOCHIRUS STIMPSON Glypturus acanthochirus Stimpson, Proc. Chicago Acad. Sci., vol. 1, p. 46, 1866; Ann. Lyc. Nat. Hist. New York, vol. 10, p. 121, 1871; de Man, Siboga- Exped., monogr. 39a", p. 19, 25, 180, 1928. De Man has already intimated that Glypturus as a genus is perhaps not distinct from Callianassa. His point is well taken, for any ex- amination of the subgenus Callichirus will show a number of species that have third maxillipeds approaching, if they do not actually possess, the Glypturus type of external maxilliped. The generic character given by Stimpson and based on these appendages, “ ischium armed along the middle of its inner [face, instead of edge as Stimpson has it—surely a slip of the pen] with a sharp, prominent, spinous crest,” is common to the majority of the species of Callianassa; as are the “deeply sculptured caudal lamellae,” to a number of the representa- tives of the subgenus Callichirus. Little needs otherwise to be said of this well-characterized and easily recognizable species. The oppor- tunity is here taken (pls. 1-4) of supplying some photographic details of a male specimen of the species. On the distal half of the inner face of the ischium of the third maxilliped (pl. 3, fig. 4) is a comb of Io spinous teeth of irregular size, but roughly, more or less alternately large and small; near the proximal margin of the joint and continuing the line of the comb are four large and one much smaller dentiform tubercles. EXPLANATION OF PLATES PLATE I Fic. Callianassa jamaicense, 3, carapace and frontal appendages, about 23. C. jamaicense var. louisianensis, 3, same. C. islagrande, 3, same. C. longiventris, 3, same. C. rathbunae, 3, same. C. acanthochirus, 3, same. ONCE CoN PLATE 2 Fic. Callianassa islagrande, 3, chelipeds, * #. C. rathbunae, 3d, same. C. longiventris, 3, same. C. jamaicense var. louisianensis, 3, same. C. acanthochirus, &, same. C. jamaicense, 3, same. C. jamaicense var. louisianensis, 3. a, d, inner and outer faces of third maxilliped ; b, c, inner and outer aspects of third leg, & about 23. C. jamaicense, 3. a, d, inner and outer faces of third maxilliped; b, c, inner and outer aspects of third leg, about 23. at ee Ss aS) be ee) NO. 2 MUD SHRIMPS—SCHMITT 21 PLATE 3 Fic. 1. Callianassa rathbunae, 3. a, d, outer and inner aspects of third leg; b, c, outer and inner faces of third maxilliped, & about 24. 2. C. islagrande, 3. a, d, inner and outer faces of third maxilliped; b, c, inner and outer aspects of third leg, & about 2+. 3. C. longiventris, 3. a, d, inner and outer faces of third maxilliped; b, c, inner and outer aspects of third leg, & about 21. 4. C. acanthochirus, 3. a, d, inner and outer faces of third maxilliped; b, c, inner and outer aspects of third leg, & about 2t. PLATE 4 Fic. Callianassa jamaicense, 3, telson, dorsal view X about 2}. . rathbunae, 3, same. . longiventris, 3, same. . jamaicense var. louisianensis, 3, same. . islagrande, 3, same. . acanthochirus, 3, same. AwnbBW ND PD AVG Dy SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR IS NOZ 2 eed MUD SHRIMPS OF THE GENUS CALLIANASSA (For explanation, see page 20.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE Ic) NOW2 ren MUD SHRIMPS OF THE GENUS CALLIANASSA (For explanation, see page 20.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93, NO. 2, PL. 3 MUD SHRIMPS OF THE GENUS CALLIANASSA (For explanation, see page 21.) (‘12 a8ed 99s ‘uorjeurldxea 410.7) VSSYNVIT1IVD SNNADSD AHL AO SDWIYHS GNW ¢ “ld ‘2 “ON ‘€6 “10A SNOILOS11090 SNOSNVIISOSIN NVINOSHLIWS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 3 NEW EARTHWORMS FROM CHINA, WITH NOTES ON THE SYNONYMY OF SOME CHINESE SPECIES OF DRAWIDA AND PHERETIMA BY G. E. GATES Judson College, Rangoon, Burma (PUBLICATION 3293) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION FEBRUARY 27, 1935 The Lord Battimore Press BALTIMORE, MD., U. & As NEW EARTHWORMS FROM CHINA, WITH NOTES ON Pot SYNONYM OF SOME CHINESE SPECIES OF DRAWIDA AND PHERETIMA ByiGs Eee GAMES Judson College, Rangoon, Burma Seven years and more ago the author undertook a study of the earthworms collected in the province of Szechuan, China, by Dr. D. C. Graham for the United States National Museum. As many of the Chinese species were inadequately characterized, completion of the report on this study had to be delayed until the gaps in our knowledge of the older forms could be filled out. In the meantime further col- lections made by Dr. Graham necessitated numerous changes in the earlier manuscript. Just recently opportunities have arisen to examine the types of many important species. As the types of the few remain- ing species are not likely to be available for examination in the im- mediate future, the report was brought to completion. As publica- tion of the full report may be delayed for some time, owing to finan- cial conditions, it seems desirable to publish diagnoses of the new species together with short notes on the changes in the synonymy. Full descriptions of the types of the new species, together with some account of the examinations of the older types, as well as explana- tions of the changes in the synonymy will be included in the definitive report. In connection with a few of the species the opportunity has been taken to include brief remarks on a very recent paper by Chen (1933), published after the writer’s longer paper was completed. DRAWIDA Michaelsen The clitellum, in this genus, appears to be a rather evanescent struc- ture, often absent in most, if not all, of the specimens submitted to the systematist for examination, and possibly present or recognizable externally during only a small portion of the year. In these circum- stances many of the species in this genus have been erected on aclitel- late specimens. Unfortunately, some of the types have been so juvenile that dis- tinguishing specific characteristics are entirely unrecognizable, and SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 3 to SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 other types are so immature that there is doubt as to whether the structures of most importance for taxonomic purposes have reached a definitive stage in their development. Criteria for the recognition of sexual maturity in the genus Drawida are therefore essential. Size (of the specimen) is not a satisfactory criterion: a worm 30 mm long may be fully mature, whereas other. worms well over 100 mm in length may be too small to identify. Other criteria that might be suggested have proved unsatisfactory, but for most taxonomic purposes sexual maturity may be regarded as evidenced by the distension of ovarian segments or chambers and ovisacs by free ova, the distension of the testis sacs by testicular material, and the distension of the spermathecal ampullae by a floc- culent, whitish material. Especial attention is necessary, in identifications and specific de- scriptions, to the characteristics and contents of invaginations at the ectal ends of the male deferent apparatus and of the spermathecae, to the central body of the prostate, to the spermathecal atria, and to the glands of the genital markings and the male porophores. The extent of the closing off of the ovarian segment is also to be determined. DRAWIDA GISTI Michaelsen, 1931 Distinguished from D. hehoensis Stephenson, 1924, by the greater size of the prostates and their relation to the penis pouches, greater length of the penes, larger size of the spermathecal atria, incomplete- ness of closure of the ovarian chamber, presence of an “ urn-shaped ” gland in the atrial wall, and by the definite genital markings with their glands projecting through the parietes into the coelom. D. gisti f{. typica Chen, 1933, differs from the types as follows: apertures of the penial chambers located in bc nearer to c than B, prostates shorter, 4 to 8 mm in length (including duct) rather than g to II mm, presence of a genital marking in the penis pouch. These differences may or may not be significant. D. gisti var. nanchangiana Chen, 1933, differs from the types as follows: prostates smaller (2 to 24 mm in length), relation of pros- tates to the penial chambers, minute size of the spermathecal atria, and “ventrally closer and more prominent setae”. According to Chen the spermathecal atrium is “ often minute, bulb-like embedded in body wall, rarely elongate like other varieties”. Specimens with minute atria embedded in the body wall may, of course, be juvenile, but if sexually mature must be specifically distinct from forms with elongate and coelomic spermathecal atria. NO. 3 NEW EARTHWORMS FROM CHINA—GATES 3 D. gisti var. anchingiana Chen, 1933, is not adequately described but differs from the types as follows: limitation of the penis pouches (?) to the body wall, small size and smooth surface of the prostates, and the presence of the spermathecal atria in vii rather than viii. These differences are important enough to distinguish the worms specifically from D. gisti. The specimens may not be sexually ma- ture—vide absence of granulations on the prostates, the empty ovisacs, and the small size of the spermathecal ampullae, as well as the indis- tinctness of the clitellum. DRAWIDA GRAHAMI, n. sp. Length 55 mm, diameter 4 mm. Spermathecal pores in 7/8 midway between b and c. Male pores in bc, nearer to 0 than to c, on small porophores seated on 10/11, intersegmental furrow 10/11 ending blindly against the lateral and median sides of the porophores. Genital markings on vii-xili, each marking with a firm, rounded gland pro- jecting through the parietes into the coelom. Vas deferens short. Prostates sessile on the parietes, outline circu- lar; central body tiny, ovoidal, pointed end buried in the parietes. Segment xi reduced to a horseshoe-shaped ovarian chamber. Sper- mathecal atria finger-shaped, erect in viii. ° Type —vU.S.N.M. no. 20093, from Suifu, Szechuan. Distinguished from D. japonica by the absence of the appendix on the ovisacs, the more median location of the spermathecal pores, and the sessile prostates. DRAWIDA JAPONICA Michaelsen, 1892 This species is characterized by a posterior continuation of the ovisac into a very long and slender rodlike appendix, which may extend as far back as segment xliii. These appendices were over- looked by both Michaelsen and Stephenson. The central body of the prostate is shortly tubular, finger-shaped. The vas deferens passes ectally into the body wall just median to the prostate. The ovarian chamber is horseshoe-shaped. The male pores are located on segment x on the ventral faces of porophores in be, nearer to b than to c. D. japonica {. siemssen Michaelsen, 1910 and 1931, is probably specifically distinct from D. japonica. It differs from the latter in size, number of gizzards, external genital markings and characteris- tics of the male porophores. Unfortunately, the internal organs of the single specimen have been lost so that the species cannot be ade- quately characterized. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 D. japonica Chen, 1933, is probably only in part, if at all, con- specific with D. japonica. Chen’s forms are characterized by greater size (70 to 200 by 3 to 54 mm rather than 28 to 65 by 1 to 3 mm), location of the male pores, at least in part, on penes or penislike pro- tuberances, and the absence of the characteristic appendices on the ovisacs. Possibly Chen’s material is to be referred to two distinct species—mention is made of “some smaller ones about 30 mm long bf and 2 mm wide also showing sexually mature ”. DRAWIDA LINHAIENSIS Chen, 1933 The types of this species are probably not fully mature—vide smooth or finely-granular surface of the prostates, the small size of the spermathecal ampullae, and the lack of clitellar development (“clitellum traceable only in one specimen”). The male porophore may possibly contain a spheroidal gland as in D. nepalensis. Further information is needed on the penis (?) pouch (?) and rela- tion of the prostate thereto, as well as on characteristics of the sper- mathecal atria and ovarian chamber in fully sexual specimens. DRAWIDA SINICA Chen, 1933 Apparently a good species—the prostates and the spermathecal atria certainly appear to be quite characteristic. The types are not, how- ever, fully mature—vide absence of ova in the ovisacs and ovarian segment and the small size and emptiness of the spermathecal ampullae. DRAWIDA SYRINGA Chen, 1933 The description is rather abbreviated and the species cannot be ade- quately characterized in absence of information with regard to the central body of the prostates and the relation of the prostatic duct to the penis pouch(?). Length of ovisacs, in view of rather wide intra- specific variation in this characteristic, may possibly be of little importance. PHERETIMA Kinberg In this genus especial attention is necessary, in identifications and specific descriptions, to the characteristics and contents of the invagi- nations at the ectal ends of the male deferent apparatus and of the spermathecae, to the intestinal caeca, to the testis sacs and their relation to the seminal vesicles and to the glands of the genital markings. The failure to describe adequately some of these characteristics has NO. 3 NEW EARTHWORMS FROM CHINA—GATES 5 been responsible for the erection of unnecessary species and for con- siderable confusion in the synonymy. Abnormalities probably occur much more frequently in this genus than in most other genera of earthworms and are often of such a nature as to render a particular specimen scarcely recognizable or indeed actually unrecognizable. Considerable taxonomic caution may therefore be necessary not only in connection with unique specimens but even with whole series or large batches. PHERETIMA ABDITA, n. sp. Length 80 to 140 mm, diameter 34 to 6 mm. .Setae: vi/30-42, vii/30-44, Xvi1/14-16, xviii/13-17, xix/16-19, xx/54-69. First dorsal pore in 12/13. Spermathecal pores small, three pairs, in 5/6-7/8. Sis Fic. 1.—Pheretima abdita, new species. Spermathecal diverticulum and portion of spermathecal duct, cleared in lactophenol. X ca. 94. Male pores at ventral ends of tubular penes located in eversible, deep parietal invaginations with elongately slitlike apertures. Genital mark- ings paired, presetal on xviii and xix. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Septa 8/9-9/10 present and thickly muscular. Intestinal caeca simple. Testis sac of x annular. Testis sac of xi U-shaped or annu- lar. Seminal vesicles of xi included in the posterior testis sac. Sper- mathecal diverticulum with a short, muscular stalk, a middle portion more or less regularly bent back and forth in a zigzag fashion, and a terminal, ovoidal seminal chamber. Genital marking glands sessile, sometimes slightly protuberant into the coelom. Type—uvU.S.N.M. no. 20094; from Suifu, Szechuan. Distinguished from P. flowert and P. gemella as well as from 'P. rockefelleri by the presence and muscularity of septa 8/9-9/10. PHERETIMA ANTEFIXA, n. sp. Length 85 to 120 mm, diameter 34 to 5 mm. Setae: viii/12-14, xiv-xv/+ or —, xvi/7-16, xvii/12-18, xvili/6-10, xix/13-17, xx/36- 42. First dorsal pore in 12/13. Spermathecal pores superficial, one pair, in 8/9. Male pores superficial, toward lateral margins of short, transverse ridges. Genital markings unpaired, median, presetal on ili, Iv, and v. Fic. 2—Pheretima antefixa, new species. Portion of spermathecal diverticulum, cleared in lactophenol. Intestinal caeca simple. Testis sacs of x and xi unpaired and ven- tral. Spermathecal diverticulum with a short, muscular stalk and an elongately tubular seminal chamber, the latter nearly straight, twisted or looped. Genital marking glands stalked and coelomic. Type—vU.S.N.M. no. 20095, from Suifu, Szechuan. Distinguished from other bithecal species of Pheretima with sper- mathecal pores in 8/9 by the extreme anterior location of the unpaired, median genital markings. NO. 3 NEW EARTHWORMS FROM CHINA—GATES 7 PHERETIMA ASPERGILLUM (E. Perrier), 1872 In the synonymy of this species there must now be placed the fol- lowing: Pheretima lauta Ude, 1905, Pheretima paraglandularis Fang, 1929, Pheretima siemssem Michaelsen, 1931 (in part). PHERETIMA BUCCULENTA, n. sp. Length 135 mm, diameter 6 mm. Setae: vi/22, vii/22, vili/25, xvii/16, xviii/20, x1ix/20. First dorsal pore in 12/13. Spermathecal pores minute and superficial, four pairs, in 5/6-8/9. Male pores on tiny, conical tubercles in the dorsalmost portions of deep parietal invaginations with longitudinally slitlike apertures. Genital markings paired, presetal on xviil. Intestinal caeca simple. Testis sacs of x and xi unpaired and ventral. Genital marking glands sessile but protuberant through the parietes into the coelom. Type—vU.S.N.M. no. 20096, from “Szechuan ”’. Distinguished from other octothecal Chinese species of Pheretima by the combination of superficial spermathecal pores and deeply invagi- nate male pores. PHERETIMA EXILIS, n. sp. Length 68 to 85 mm, diameter 2 to 24 mm. Setae: vi/39, xvi/6, Xvii/10-15, xvili/8, xix/II-13, xx/50. First dorsal pore in 12/13. Spermathecal pores minute and superficial, two pairs, in 5/6-6/7 or on the posteriormost margins of v and vi. Male pores within slight depressions on glistening, not sharply demarcated areas. Genital mark- ings paired on xvii and xix, probably postsetal. Intestinal caeca simple. One pair of testis sacs in x. One pair of vertical testis sacs in xi. Seminal vesicles of xi included within the posterior testis sacs. Genital marking glands sessile but slightly pro- tuberant through the parietes into the coelom. Type.—uU.S.N.M. no. 20097, from Suifu, Szechuan. Distinguished from quadrithecal Chinese species of Pheretima with spermathecal pores in 5/6-6/7 by the inclusion of the seminal vesicles of xi within the posterior testis sacs. PHERETIMA FLEXILIS, n. sp. Length 40 mm, diameter 2 mm. Setae; vii/16, viii/16, xvii/I10, XVili/10, xix/1I. First dorsal pore in 13/14. Spermathecal pores minute and superficial, three pairs, in 6/7-8/9. Male pores superficial, at the centers of tiny, transversely oval areas, each area surrounded by 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Fic. 3—Pheretima exilis, new species. Spermatheca with iridescent mass in seminal chamber. X ca. 64. Fic. 4.——Pheretima exilis, new species. Spermatheca with no iridescence in diverticulum. X ca. 64. Fic. 5.—Pheretima fornicata, new species. An ectal portion of spermathecal diverticulum, cleared in lactophenol. Fic. 6.—Pheretima fornicata, new species. Ental portion of spermathecal di- verticulum, cleared in lactophenol. Magnification much less than in fig. 5. NO. 3 NEW EARTHWORMS FROM CHINA—GATES 10) several concentric furrows. Genital markings unpaired and median, presetal on viii, postsetal on xvii and xviii. Intestinal caeca simple. Testis sac of x horseshoe-shaped. One pair of vertical testis sacs in xi. Seminal vesicles of xi included within the posterior testis sacs. Spermathecal diverticulum with short, muscular stalk and an elongately tubular seminal chamber, the latter variously bent, twisted or looped. Genital marking glands with long, coelomic stalks. Type—uU.S.N.M. no. 20098, from between Gin Keo Ho and Dawei, Szechuan. Distinguished from P. hupeiensis by the absence of septa 8/9-9/10 and from P. lohri by characteristics of the testis sacs as well as by the extra pair of spermathecae. PHERETIMA FORNICATA, n. sp. Length 78 to 90 mm, diameter 4 to6 mm. Setae: vi/17-24, vii/19- 21, vili/18-23, xvii/13-14, Xvil1/Q-14, Xix/12-15, xx /56; a wide dorsal gap in the setal circle of ii. First dorsal pore in 12/13. Spermathecal pores minute and superficial, four pairs, in 5/6-8/9. Male pores superficial, on circular to transversely oval, disk-shaped porophores. No genital markings. Septum 8/g present but membranous. Intestinal caeca simple. Testis sacs of x and xi unpaired and horseshoe-shaped. Spermathecal diverticulum with a long, slender stalk and a spheroidal or asymmetri- cal seminal chamber. Type —uvU.S.N.M. no. 20099, from Tatsienlu, Tibet. Distinguished from P. hongkongensis by the gap in the setal circle of ii, the exclusion of the seminal vesicles of xi from the posterior testis sac, and the absence of genital markings. PHERETIMA GRAHAMI, n. sp. Length 235 to 285 mm, diameter II to 15 mm. Setae: vii/22-25, vili/22-27, xvii/26, xvili/19, xix/19-25, xx/80-91. First dorsal pore in 12/13-13/14. Spermathecal pores on tiny, conical protuberances into large, club-shaped spermathecal chambers, the latter invaginated posteriorly and deeply into the coelom and bound by connective tissue to the ventral parietes; apertures of spermathecal chambers large, transversely slitlike, three pairs, in 6/7-8/9. Male pores on broadly conical tubercles in large copulatory chambers with apertures approxi- mating to transversely slitlike. External genital markings lacking; IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 5 to 6 circular to oval, flat-surfaced markings in each copulatory chamber; one large, oval marking on the dorsal (morphologically anterior ) wall of each spermathecal chamber. Intestinal caeca simple. Testis sacs of x and xi unpaired and ven- tral. Spermathecal diverticulum with a short, muscular stalk and an Fic. 7—Pheretima grahami, new species. Portion of spermathecal duct, cleared in lactophenol. elongately tubular seminal chamber, the latter often looped in a regu- larly zigzag fashion. Type—uvU.S.N.M. no. 20100, from Da Shiang Lin Pass, Szechuan. Distinguished from P. vulgaris by the unpaired, ventral testis sacs and the large, club-shaped spermathecal chambers with their large, oval genital markings. PHERETIMA GUILLELMI (Michaelsen), 1895 A good species. Distinguished from P. houlleti (E. Perrier), 1872, with which it has been confused, by the limitation of the male pore invaginations to the parietes (in P. houlleti the male pores are in copu- latory chambers). P. houlleti Stephenson, 1925, and P. ichangensis Fang, 1933, must be included in the synonymy. Amyntas houlleti Mi- chaelsen, 1899, is probably also a synonym. P. vulgaris agricola Chen, 1930, is probably also, at least in part, a synonym. P. guillelmi Chen, 1933, is a composite of P. guillelmi and P. vulgaris. PHERETIMA HONGKONGENSIS Michaelsen, 1910 The unpaired testis sac of x is ventral and median. The unpaired testis sac of xi is U-shaped, the limbs of the U reaching to the dorsal blood vessel. The seminal vesicles of xi are included within the posterior testis sac and are surrounded by a layer of testicular coagulum. The holotype is either not fully sexual or is not quite normal. NO. 3 NEW EARTHWORMS FROM CHINA—GATES Ii PHERETIMA HUPEIENSIS (Michaelsen), 1895 The testis sacs of x and xi are U-shaped. Chen (1933) appears to regard the testis sacs as annular. Chen’s account is not quite clear. He places the seminal vesicles of xii in a membranous sac similar to that which contains the seminal vesicles of x1, though there is no testis sac in xii. Coelomic coagulum may sometimes be compacted around the seminal vesicles so closely as to produce an appearance somewhat similar to that of a testis sac filled with testicular coagulum. Perichaeta hupeiensis Gee, Boring, and Wu, 1927, is certainly not P. hupetensis and is probably a composite of three distinct species. PHERETIMA IGNOBILIS, n. sp. Length 55 mm, diameter 3 mm. Setae: vi/1I7, vii/16, viii/16, Xvli/I5, xvili/9Q, x1x/16. First dorsal pore in 11/12. Spermathecal pores in parietal invaginations with transversely slitlike apertures, four pairs, in 5/6-8/9. Male pores on the roof and toward the median side of transversely slitlike depressions. No genital markings. Intestinal caeca simple. Testis sacs of x and xi paired and ventral. Type—vU.S.N.M. no. 20101, from near Ningyuenfu, Szechuan. Distinguished from all octothecal Chinese species of Pheretima by the location of the spermathecal pores in parietal (or deeper?) invaginations. PHERETIMA LIMELLA, n. sp. Length 60-85 mm, diameter 24-5 mm. Setae: v/38, vi/51, xiv- xvi/ +, xvli/18-29, xviii/14-21, xix/18-26. First dorsal pore in 12/13. Spermathecal pores minute and superficial, one pair, in 5/6. Male Fic. 8—Pheretima limella, new species. Spermathecal diverticulum, cleared in lactophenol. pores at centers of small, oval tubercles; a thin fold of tissue at the lateral margin of each male porophore can be drawn mesially over the tubercle in the manner of an eyelid. Genital markings paired and presetal on xvii. 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Septa 8/9-9/10 present and thickly muscular. Intestinal caeca simple. Testis sacs of x and xi unpaired. Spermathecal diverticulum with a muscular stalk and an elongate seminal chamber, the ectal portion of the latter looped, the ental portion ovoidal. Genital marking glands sessile. Type—vU.S.N.M. no. 20102, Suifu, Szechuan. Distinguished from P. zoysiae Chen, 1933, by the presence and muscularity of septa 8/9-9/10 and by the presence of genital markings. PHERETIMA MIRABILIS (Bourne), 1887 In the synonymy of this species there must now be placed P. hetero- chaeta (Michaelsen), 1891, and P. divergens var. yunnanensis Ste- phenson, 1912. PHERETIMA MODESTA Michaelsen, 1927 The male pores are in copulatory chambers which, in a completely retracted condition, protrude rather conspicuously into the coelom. A very large portion of the coelomic protuberance is composed of tiny quirks in the prostatic duct on the dorsal face of the chamber and a mass of connective tissue surrounding these quirks. The lumen of the chamber is rather small but is narrowed in the outer layers of the parietes as if by a contraction of an annular, sphincter muscle. In slightly softened specimens the apertures of the chambers gape open, and the lumen of the chamber does not extend internally beyond the level of the coelomic face of the parietes. In the synonymy of P. modesta are to be placed P. kiangensts Michaelsen (in part only?) and P. hesperidum Chen, 1931 and 1933. The status of Beddard’s P. hesperidum was discussed in Gates (1932). Stephenson (1933) considered the status of P. hesperidum and has also refused to accept Beddard’s species. PHERETIMA OBSCURITOPORA Chen, 1931 Erected on juvenile specimens of uncertain age, in which the adult and hence definitive specific characteristics are unrecognizable. The Szechuan specimens might be rather small juveniles of P. grahanu or P. praepinguis; the Nanking specimens may possibly belong to P. tschiliensis. PHERETIMA OMEIMONTIS Chen, 1931 P. paraglandularis var. omeimontis Chen, 1931, is distinguished from P. paraglandularis Fang, 1929 (= P. aspergillum), by differ- ences so numerous and significant that omeinontis must be regarded as specifically distinct. NO. 3 NEW EARTHWORMS FROM CHINA—GATES 13 PHERETIMA PAETA, n. sp. Length 75-136 mm, diameter 5-6 mm. Setae: viii/22-24, xvii/18-21, Xvili/10-16, xix/21-24, xx/65-68. First dorsal pore in 11/12-12/13. Spermathecal pores on circular to oval areas within deep invagina- tions with transversely slitlike apertures, two pairs, in 7/8-8/9. Male pores on the dorsal wall of large copulatory chambers conspicuously protuberant into the coelom. On the median wall of the copulatory chamber a presetal, transversely oval genital marking, occasionally also a postsetal marking ; on the roof of the chamber one or two further markings of variable shape and:size. External genital markings paired, on the posteriormost margins of vii and viii, each marking one to three intersetal intervals median to the aperture of the spermathecal invagination. Intestinal caeca compound, dorsalmost secondary caecum the longest. Testis sacs of x and xi unpaired and ventral. Spermathecal diver- ticulum with a muscular stalk and an elongately tubular seminal cham- ber, the latter looped back and forth in a regularly zigzag fashion. Type —vU.S.N.M. no. 20103, from Song Pan, Szechuan. Distinguished from P. omeimontis by the copulatory chambers, and from P. schmardae by the larger size, the invagination of the sper- mathecal pores, and the presence of genital markings. PHERETIMA PAPILLIFERA, n. sp. Length 100 mm, diameter 4 mm. Setae: vi/11(+ ?), vii/13(+ ?), sally on iv. First dorsal pore in 11/12. Spermathecal pores minute and superficial, three pairs, in 5/6-7/8. Male pores superficial, at centers of circular areas demarcated by slight furrows. Genital markings paired, presetal, on xi-xiv. Intestinal caeca simple. Testis sacs of x and xi paired. Spermathe- cal diverticulum with a short muscular stalk and a longer, more irregu- lar seminal chamber. Genital marking glands sessile on the parietes. Type—vU.S.N.M. no. 20104, from near Zachoo, Szechuan. Distinguished from sexthecal Chinese species of Pheretima with dorsally on iv and by the location of the genital markings. PHERETIMA PECTENIFERA Michaelsen, 1931 The male pore region of P. pectenifera is so remarkably similar to that figured for P. yamadai Hatai, 1930, that the former may be, in reality, a synonym of the latter. Hatai’s species is not adequately 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 characterized and P. pectenifera must, accordingly, be allowed to stand until after reexamination of the Japanese types or amplification of the original description. Further information is needed with regard to size, spermathecal pores, setal numbers, septa 8/9-9/10, intestinal caeca, testis sacs, and the preclitellar genital markings. Pheretima yamadai Chen, 1933, is almost certainly a composite of two distinct species, P. pectenifera and P. tschiliensis Michaelsen, 1928. Chen himself distinguishes two forms, A and B; A is probably tschiliensis, B probably pectenifera. P. pingi Michaelsen, 1931, may be, in part, synonymous with P. pectenifera. At least four of the Hamburg specimens labelled “P, pingi”’ are obviously P. pectenifera. PHERETIMA PINGI Stephenson, 1925 The hearts of x are present but are small, usually empty and bound by connective tissue to the anterior face of 10/11. In these circum- stances the hearts of x are easily overlooked. According to Chen (1933) the hearts of x are entirely lacking. Fic. 9.—Pheretima pingi. An ental portion of spermathecal diverticulum, cleared in lactophenol. (The term “heart” is used to refer to a segmental commissure connecting the ventral blood vessel with the supra-esophageal vessel or the dorsal blood vessel or both, in any of segments ix-xi11.) PHERETIMA POMELLA, n. sp. Length 87 mm, diameter 5 mm. Setae: vii/19, xvii/19, xvili/14, x1x/19, xx/ca. 50; lacking dorsally on ii. First dorsal pore in 10/11. Spermathecal pores minute and superficial, two pairs, presetal on vii and viii, slightly nearer to the intersegmental furrows than the setal circles. Male pores superficial, on rather indistinctly demarcated areas. Genital markings paired, presetal on xii, xiii and xvili, post- NO. 3 NEW EARTHWORMS FROM CHINA—GATES I5 setal on xviii; markings on xviii in line with male pores, on xii-xiii about in ab. Intestinal caeca simple. Testis sacs of x and xi unpaired and ventral. Spermathecal diverticulum with a short, muscular stalk and an elon- gately tubular seminal chamber, the latter twisted into a ball-like mass of loops. Type.—vU.S.N.M. no. 20105, from Suifu, Szechuan. Distinguished from P. planata Gates, 1926, by the posterior loca- tion of the spermathecal pores, the absence of copulatory chambers, and the locations of the genital markings. PHERETIMA PRAEPINGUIS, n. sp. Length 207-(357?) mm, diameter 16 mm. Setae: vii/23, viii/24, Xvli/20, xvili/Q+, xix/22, xx/93. First dorsal pore in 12/13. Sper- mathecal pores on tiny tubercles located in parietal invaginations with transversely slitlike apertures, three pairs, in 6/7-8/9. One circular genital marking on the anterior wall of each invagination. Male pores on tubercles in the lateralmost portions of deep parietal invaginations with crescentic apertures, lateral walls of the invaginations thin and nonsetigerous. Just median to each male pore tubercle a single genital marking ; on the median wall of the male invagination a transversely oval, presetal genital marking. External genital markings paired, presetal on vii, vili and ix. Intestinal caeca simple. Testis sacs of x and xi unpaired and ventral. Spermathecal diverticulum with a short, muscular stalk and an elon- gately tubular seminal chamber, the latter with several slight constrictions. Type—vU.S.N.M. no. 20106, from Mount Omei, Szechuan. Distinguished from P. tschiliensis by the spermathecal invaginations and the genital markings therein. PHERETIMA ROBUSTA (E. Perrier), 1872 In the synonymy of this species there must now be placed the fol- lowing: P. siemssent Michaelsen, 1931° (in part), P. fokiensis Michaelsen, 1931, P. lauta Ude, 1932. P. léhri (Michaelsen), 1899, can be distinguished from P. robusta only by the small size. The types of P. lohri may be dwarfed forms of P. robusta, the dwarfing the result of a heavy infection of parasitic protozoa. P. lauta Chen, 1933, is probably in large part, if not entirely, synonymous with P. robusta. Chen does, however, differentiate between ‘“‘ coast”? and “ inland” forms, and possibly some of one or both groups may be referable to 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 P. aspergillum. Chen’s specimens appear to be more or less ab- normal—* prostates not well developed... . or totally absent or very rudimentary ” Fic. 10.—Pheretima robusta. Portion of a spermatheca, cleared in lactophenol. X ca. 54. All Szechuan specimens that can be referred to P. corrugata Chen, 1931, are abnormal. The two paratypes that have been available for examination are also abnormal. P. corrugata is therefore regarded as very dubious, possibly synonymous with P. robusta. Fic. 11.—Pheretima schmardae. Duct and diverticulum of a spermatheca. Cd 70: PHERETIMA TSCHILIENSIS MICHAELSEN, 1928 In the synonymy of this form there must now be placed the fol- lowing: P. asiatica Michaelsen, 1902 (in part), P. tibetana Michael- sen, 1931, and P. kiangsuensis Chen, 1930 and 1931. NO. 3 NEW EARTHWORMS FROM CHINA—GATES 17, Fig. 12—Pheretima ssechuanensis. Spermathecal duct and diverticulum, cleared in lactophenol. X ca. 52. Fic. 13—Pheretima tschiliensis. Spermathecal duct and diverticular stalk, cleared in lactophenol. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Types of P. asiatica (Michaelsen), 1900, have not, unfortunately, been available for examination, but inasmuch as Michaelsen’s reasons for splitting off P. tibetana from P. asiatica do not appear to be ade- quate—especially in view of the variability of the form under con- sideration—it is quite possible that P. tschiliensis may have to be placed in the synonymy of P. asiatica. The latter is not, however, sufficiently distinguished from P. guillelmi to warrant further change at the present. Fic. 14.—Pheretima tschiliensis. Spermathecal diverticulum, cleared in lacto- phenol. X ca. 45. PHERETIMA TUBERCULATA, n. sp. Length 80-110 mm, diameter 3-5 mm. Setae; vi/g-10, vii/9-14, Xvli/16-19, Xvili/10-13, xix/17-20, xx/40-53. First dorsal pore in 10/11. Spermathecal pores minute and superficial, three pairs, in 5/6- 7/8. Male pores superficial, at centers of small, oval areas. Genital markings small tubercles in immediate vicinity of male and spermathe- cal pores. Intestinal caeca compound, dorsalmost secondary caecum the long- est. Testis sacs of x and xi unpaired and ventral. Spermathecal diverticulum with a muscular stalk and an elongately tubular seminal chamber, the latter often looped in a regularly zigzag fashion. Genital marking glands stalked and coelomic, stalks erect in the coelom. Type—vU.S.N.M. no. 20107, from Suifu, Szechuan. Distinguished from sexthecal Chinese species of Pheretima with spermathecal pores in 5/6-7/8 by the compound intestinal caeca. NO. 3 NEW EARTHWORMS FROM CHINA—GATES 19 PHERETIMA VULGARIS Chen, 1930 A good species if the account of the typical forms is correctly inter- preted, distinguished from P. guillelmi, with which Chen has lately (1933), confused it, by the characteristic club-shaped copulatory chambers as well as by the spermathecal invaginations. Fic. 15.—Pheretima vulgaris. Spermathecal duct and ectal portion of divertic- ulum. X ca. 36. P. pingi Michaelsen, 1931, and P. kiangensis Michaelsen, 1931, are probably, at least in part, synonymous with P. vulgaris; some of the Hamburg specimens labelled P. pingi and P. kiangensis are obviously P. vulgaris. An aclitellate specimen from Ichang in Hupeh, referred by Fang to P. vulgaris, is correctly identified, but specimens from Peiping referred to P. vulgaris by Fang cannot be properly placed. LITERATURE CITED CuHen, Y. 1933. A preliminary survey of the earthworms of the lower Yangtze valley. Contr. Biol. Lab. Sci. Soc. China, vol. 9, no. 6. Gates, G. E. 1932. The earthworms of Burma. III. The Megascolecinae. Rec. Indian Mus., vol. 34, pt. 4. STEPHENSON, J. 1933. Oligochaeta from Australia, North Carolina, and other parts of the world. Proc. Zool. Soc. London, 1932. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 4 Peo NEER: WIND) TUNNELS (With 4 PLateEs) BY N. H. RANDERS-PEHRSON Assistant, Division of Aeronautics, Library of Congress (PUBLICATION 3294) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JANUARY 19, 1935 The Lord Battimore Press BALTIMORE, MD., U. 8. A. PIONEER WIND TUNNELS By N. H. RANDERS-PEHRSON Assistant, Division of Aeronautics, Library of Congress (WitH 4 PLatTEs) In a paper read before the Royal Society in 1759 John Smeaton presented the following very neat outline of the various methods that may be used for aerodynamic research: * In trying experiments on wind mill sails, the wind itself is too uncertain to answer the purpose; we must have recourse to an artificial wind. This may be done two ways; either by causing the air to move against the machine, or the machine to move against the air. To cause the air to move against the machine, in a sufficient volume, with steadiness and the requisite velocity, is not easily put in practice: To carry the machine forward in a right line against the air, would require a larger room than I could conveniently meet with. What I found most practicable, therefore, was to carry the axis, whereon the sails were to be fixed, progressively round in the circumference of a large circle. Technical difficulties prevented Smeaton from using a wind tunnel and prompted him to adopt the whirling machine, invented in 1746 by Ellicott and Robins. This and other inferior methods of research were in use long after wind tunnels had been constructed and found satisfactory. Thus, Eiffel spent much time dropping plates from the tower bearing his name before adopting the wind-tunnel method ; Langley and others used whirling machines, while many used natural wind, which Smeaton had already found to be unreliable. F. H. WENHAM The distinction of being the first to introduce the wind tunnel be- longs to Francis Herbert Wenham, founder member of the Aero- nautical Society of Great Britain, who read at its opening meeting his classical paper on “ Aerial Locomotion ’’. In 1871 this Society desired to undertake systematic aerodynamic experiments to obtain “data on which a true science of aéronautics can be founded”. A subscription fund was established ; an instrument designed by Wenham *Smeaton, John, Experimental enquiry concerning the natural powers of wind and water, p. 38, London, 1794. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 4 74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 was approved by the experimental committee and was constructed by John Browning, an optician and member of the Society. It was set up at Messrs. Penn’s Marine Engineering Works at Greenwich, where the world’s first wind tunnel experiments took place. The tunnel was a wooden trunk 18 inches square and 10 feet long. Through it was directed the blast from a fan, driven by a steam en- gine. The wind velocity was measured with a water gauge, various speeds up to 40 miles per hour being used. The wind was not steady, considerable fluctuations making the observations difficult. The di- rection of the wind was tested with a vane and said to be fairly straight, although there is no mention of a wind straightener of any kind. The balance was exhibited and explained to the Society by Mr. Wenham. It consisted of a vertical steel spindle, supported on a hardened steel center. Through an eye at the upper end of the spindle passed a horizontal weighing beam, supported by a cross pin axle. The long end of the beam carried the testing planes which could be set at various angles of incidence while they were always kept at right angles across the current. The short end carried a sliding counterweight so as to balance the testing plane. The drag was measured by a spring steelyard connected toa lever from the vertical spindle, close to the base of the machine. The lift was read off by a vertical spring steel- yard. The balance with the testing planes was placed in front of the tunnel at a distance of 2 feet, a wooden shield covering the balance and leaving only the planes exposed to the wind. Lift and drag were measured simultaneously, two persons making the observations. Only plane surfaces were tested, the largest being 18 inches across, the same width as the tunnel. They were placed at various angles from 15° to 60°; tests on smaller angles were found to be very desirable but could not be achieved with the instrument at hand. In spite of the crudeness of the tunnel and the shortness of the time allotted for experiments, the results were the most satisfactory of the kind obtained to that time. The experiments were very encouraging to aviation enthusiasts, as they proved that the lift at small angles exceeds the drag to a much greater extent than had previously been suspected. The desirability of a large aspect ratio and the location of the center of pressure near the leading edge were also demonstrated. The test data were published in tabular form in the Report of the Aeronautical Society. These tables were widely used and were also made the basis for actual construction, particularly by Thomas Moy NO. 4 PIONEER WIND TUNNELS—RANDERS-PEHRSON .| 3 for his “ aerial steamer ’’, the first large power-driven airplane model to rise from the ground in tethered flight.’ Twenty-five years later Wenham expressed the wish that he might have an opportunity to build a large tunnel that would convey a current “at rates varying from a gentle breese, up to a tornado that could rip the clothes off your back, or blow you away like a feather, but no flying man should mind this effect.” In 1900, at the age of 76, he actually rigged upa fan blower for experiments ; it ran at 1,700 revolu- tions per minute and gave a current of 25 miles per hour. Apparently it was driven by hand, as he says: ° I could not get beyond this as it absorbed all my strength to work it, still the current was definite and steady with proper arrangement to measure lift and drift [i. e., drag]. I attached various models in the blast, consisting of different forms of supporting surfaces. 25 miles an hour would be a sufficient speed to begin to fly with. HORATIO PHILLIPS Next to use a wind tunnel was another Englishman, Horatio Phillips. He produced his air current by means of a steam jet, hoping in that way to avoid the fluctuations of the wind which had marred Wenham’s experiments. Section on ling oe c-) LE: : . Fo os [ai beer iz =) Fic. 1.—Phillips’ wind tunnel, 1884. Phillips’ tunnel was 17 inches square and 6 feet long. Attached to one end was “‘ an expanding delivery tube of sheet-iron”’, which was 6 feet long, 12 inches wide where it entered the box, eontmedne to 8 inches, and again expanding to 2 feet. In its narrowest part was introduced a ring of iron pipe pierced with holes, through which steam was fed from a large boiler under 70 pounds pressure. This produced by suction an air current in the square part of the tunnel. In order to increase the speed of the current, the square box was partly closed by a * Aeronautical Society of Great Britain, 6th Ann. Rep., pp. 75-78, 1871; 7th Ann. Rep., pp. 6-12, 1872; 9th Ann. Rep., pp. 6-7, 1874. * Unpublished letters from F. H. Wenham to Octave Chanute, now in the Library of Congress. 4. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 solid packing of wood, leaving a space 10 by 17 inches in cross-section, where a speed up to 60 feet per second was obtained, as measured by a water gage. The balance consisted of two uprights, pivoted at the base and con- nected by a horizontal wire at the top. This passed through eyes on two stiff wires attached to the leading edge of the testing surface. Drag was measured by weights in a scalepan, attached to the model mounting by a string running over a pulley, and lift was measured by a weight suspended under the testing surface at the supposed center of pressure. The balance was pushed into the tunnel where the current was swiftest, so that the scalepan was outside the tunnel, and the suspended weight in a hole in the wood packing.’ Fic. 2.—Phillips’ balance. While Wenham studied planes only, Phillips turned his attention to curved surfaces. His were the first systematic studies of cambered air- foils, a subject about which practically nothing was known before. Phillips noticed the partial vacuum above the airfoils. He patented a number of profiles, and introduced the downward curving leading edge, now in almost universal use. Several of the airfoils developed by Phillips had a maximum lift-to-drag ratio of about 10, an efficiency adequate for pioneer flying and not known to have been surpassed before the arrival of modern wind tunnels.’ The “ Venetian blind ” airplane built by Phillips, on the basis of data obtained in his wind tunnel, readily lifted itself in tethered flight, and was, with its cambered surfaces, a distinct improvement over its predecessors. LUDWIG MACH Dr. Ludwig Mach of Vienna in 1893 was the first to use a wind tunnel to photograph the flow of air. The tunnel had a cross section of 18 by 25 centimeters ; one side was of glass and the others black on the inside. The air was sucked through by means of a centrifugal fan “Engineering, vol. 40, pp. 160-161, illus., London, Aug. 14, 1885. ° British Patent, no. 13,768, 1884; and no. 13,311, 1801. INO: a! PIONEER WIND TUNNELS—RANDERS-PEHRSON 5 at the rate of 10 meters per second. A piece of wire mesh over the opening served to straighten the current. By the use of silk threads, cigarette smoke and glowing particles of iron, the flow could be ob- served. Streams of heated air were also introduced, invisible to the eye, but recording on a photographic plate. A series of good flow photographs was obtained. JOHAN IRMINGER AND H. C. VOGT The first wind tunnel measurements of pressure distribution were made by Johan Irminger, and H. C. Vogt, of Copenhagen. Vogt, who was a marine engineer, had made extensive studies on sails and air propellers and had found that the partial vacuum on the leeward side was responsible for the greater part of the thrust. Phillips was first to notice a rarefaction, but did not press his investigation of this factor very far. Vogt, in conjunction with Irminger, director of the Copen- hagen Gas Works, undertook a series of wind-tunnel experiments to establish this fact conclusively. Fic. 3.—Irminger and Vogt’s tunnel, 1894. There was at the gas works a smokestack 100 feet high and 5 feet in diameter, serving a large number of gas furnaces. In order to utilize the draft in this chimney for experimental purposes, an opening was made in its side and a rectangular box inserted, 40 inches long and 44 by 9 inches inside cross-section. The inside of the box was polished and a shutter was used to control the speed of the air current, which ranged from 24 to 48 feet per second. To determine the pressure distribution on plane surfaces, two pieces of sheet iron were placed 1/10 inch apart, joined along the edges to form a shallow closed box. To the interior of this a water gage was *Zeitschr. Luftschiffahrt und Phys. Atmosphare, vol. 15, pp. 129-139, pls. I-III, 1806. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 connected by means of a pipe. A number of small holes were made in both surfaces, of which one at a time was opened. Two such testing planes were used ; both were 14 inches wide, one 44 inches long, reach- ing entirely across the tunnel, the other 24 inches long. The testing Section x-y Fic. 4.—Pressure measuring apparatus used by Irminger and Vogt. planes were placed in the middle of the tunnel and could be set at different angles that were indicated by a pointer on the outside. Measurements of pressure distribution were also made on bodies of various shapes, as prisms, spheres, etc., and on models of buildings and gas tanks." *Ingenioren, p. 101, Copenhagen, 1804. Inst. Civil Eng., Minutes of Proc., vol. 118, pp. 468-472, 18094. Engineering, vol. 60, pp. 787-788, illus., London, Dec. 27, 1805. NO. 4 PIONEER WIND TUNNELS—RANDERS-PEHRSON aL CHARLES RENARD Col. Charles Renard, constructor of the famous airship La France, conducted a large number of aeronautical experiments at the Etablisse- ment Militaire de Chalais Meudon, of which he was the director. Renard continued his experiments during a long period of years and employed a variety of methods and equipment. The details were secret at the time, and the information available is still meager. Fic. 5.—Renard’s tunnel, 1806. Some time during the latter half of the nineties a wind tunnel was used. It was cylindrical, 80 centimeters in diameter and 4 meters long. The wind was produced by a blower fan and said to be “ violent ”’. Fourteen meters per second is given in a published chart. There is no mention of any means for straightening the wind. In his tunnel Renard studied the stability and critical speed of air- ships.” Most of his equipment—balances, testing models, etc.—are pre- served in the aeronautical museum at Chalais Meudon. SIR HIRAM MAXIM For the construction of his giant airplane, Sir Hiram Maxim realized the necessity for scientific data and utilized a number of testing devices. Among these was a wind tunnel which was in operation in 1896. Maxim’s tunnel was a wooden box 12 feet long and 3 by 3 feet inside cross-section, connected with a shorter box 4 feet square. Two air- screws on the same shaft, placed in the wider section and driven by a 100-hp. steam engine, blew the air through the tunnel. To straighten the airstream a number of wooden slats were placed in the tunnel horizontally, vertically, and diagonally. The objects to be tested were °C, R. Acad. Sci., vol. 138, p. 146, June 6, 1904. Aerophile, vol. 12, pp. 153-155, July 1904. IV congres internationale d’aeronautique, Nancy 1909. Proces-verbeaux, rapports et memoires, p. 241. Aeronautique, vol. 6, p. 84, illus., Paris, April 1924. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 placed in front of the tunnel and the balance measured both lift and drag. Sir Hiram tested airfoils, struts, and airplane parts in his wind tunnel, and also the efficiency of steam condenser pipes.’ PAUL LaCOUR Two wind tunnels were used by Paul LaCour, of Askovy, Denmark, for windmill research. Both were made of sheet iron, cylindrical, 2.2 meters long; one was 1 meter in diameter and the other 3 meter. The wind was produced by electric blower fans and straightened by radial fins inside the tunnels. A speed of 10 meters per second was used, and this was kept constant by controlling the fan speed, which was read from a tachometer. The testing surfaces and windmill models were placed 1 meter out in front of the tunnel.” ETIENNE MAREY Etienne Marey, of Paris, famous for his chronophotographic studies of animal locomotion, in 1899 turned his attention to obtaining photo- graphs of air in motion. This was achieved by the use of narrow bands of smoke in a small, vertical wind tunnel. The tunnel was 20 by 30 centimeters in cross-section, with front and sides of plate glass and the back covered with black velvet. The air was drawn down through the tunnel by a small suction fan and straightened by passing through fine silk gauze of very even weave. Smoke was supplied through a row of fine tubes at the top of the tunnel and descended in straight bands, clearly showing the flow past small models that were inserted. Photographs were taken by means of a magnesium flash, burnt in a ventilated box close to one side of the tunnel. Among the scientists that were interested in these experiments was Samuel P. Langley. He provided funds from the Smithsonian Insti- tution for their continuance, and the next year Marey built a new and improved tunnel. This was 20 by 50 centimeters in cross-section, and the smoke tubes, 60 in number, could be made to vibrate laterally 10 times a second. * The Aeronautical Annual, 1896, pp. 50-61, illus., Boston. Maxim, Hiram, Artificial and natural flight, pp. 50-61, illus., New York and London, 1908. * LaCour, Paul, Forsogsmollen, pp. 14-15, Copenhagen, 1900. Ingenioren, no. 10, Copenhagen, 1897. NO. 4 PIONEER WIND TUNNELS—RANDERS-PEHRSON 9 The speed of the air at any point was indicated by the undulations of the smoke bands caused by these vibrations. Judging from some of the photographs where the measuring rod is seen, the speed was about 30 centimeters per second. A number of very beautiful flow photographs were obtained by Dr. Marey.” A. F. ZAHM The first complete wind-tunnel laboratory, equipped for a wide range of aerodynamical experiments and with instruments capable of exact measurements, was devised by Dr. A. F. Zahm and erected on the grounds of the Catholic University of America in the winter of 1901. This laboratory was made possible by Hugo Mattullath, inventor of a giant flying boat, Dr. Zahm having agreed to become, during his spare time, the consulting engineer of Mattullath’s company. The laboratory building was a one-story frame structure 30 by 80 feet and housed a wooden tunnel 6 feet square in cross-section and 40 feet long, with windows in the ceiling and walls. The wind was drawn through at a speed of 27 miles per hour by a 5-foot suction fan, driven by a 12 hp. electric motor. The intake end was covered with one or two screens of cheese cloth or wire mesh to straighten the wind. The air speed was held constant within a fraction of 1 per- cent by a boy with a tachometer and a rheostat, controlling the fan speed. For some researches movable liners were introduced in the main tunnel, making the current contract trumpetwise to gain speed, then run straight in a narrower stream, and finally discharge as an open jet in the after part of the main tunnel. The testing model was placed either between the parallel sides, where the wind speed was greatest, or in the center of the current where it entered the experi- mental chamber. The wind tunnel was equipped with a variety of instruments in- vented by Dr. Zahm for showing the character of the air flow and its action on the models. ™C. R. Acad. Sci., vol. 131, pp. 160-163, July 16, 1900; vol. 132, pp. 1291- 1296, June 3, IgoI. Ann. Rep. Smithsonian Inst., 1901, pp. 14, 332, 337-340, pls., 1902. Journ. phys. theroique et appliquée, 4th ser., vol. 1, pp. 129-135, illus., 1902. Scientific American, n. s., vol. 86, pp. 75-76, illus., Feb. 1, 1902. Nogues, P., Recherches expérimentales de Marey sur le mouvement dans Yair. France. Min. de lair. Publ. sci. et techn., pp. 94-07, illus., 1933. IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 The air speed was measured with a pitot-static tube, connected with an extremely sensitive manometer.” This consisted of two thin metal cups, inverted over coal-oil and supported from opposite ends of a weighing beam. Two tubes, one from underneath each cup, were joined respectively to the inner and outer tubes of the speed nozzle. To test the accuracy of this instrument a “ balloon anemometer ’’ was devised. A toy balloon floating downstream intersected on its way two thin pencils of light focussed on the moving plate of a long camera constructed for that purpose. This was an adaptation of the ingenious chronograph previously invented by Dr. Zahm for his re- searches on the speed of bullets.” The manometer was also used for the study of pressure distribution. Several aerodynamic balances were developed, among them the wire suspension balance, now in general use, and the bell crank balance, now often called the N. P. L. balance. It was called the “ universal pressure balance’ in 1902, and consisted of a bell crank with hori- zontal axle mounted on knife edges above the tunnel, having a gradu- ated horizontal arm with scalepan and sliding weights, and a vertical arm running down through a streamline wind shield to hold the models in the air stream. The laboratory was built and equipped early in 1901 and a descrip- tion was communicated to the American Association for the Advance- iment of Science June 30, 1932, and was privately printed (200 copies ) in a small pamphlet which is now a great rarity.” Mattullath died in December 1902, and the flying-boat project was abandoned, but the scientific work in the laboratory went on inter- mittently until 1908. Money grants for special researches were made by the Smithsonian Institution and the Carnegie Institution in 1904 and 1905. Results of the investigations were communicated to scientific journals and societies. The most important of these was Dr. Zahm’s ” Exhibited before the Washington Philosophical Society, May 24, 1902; described in Phys. Rev., vol. 17, pp. 410-423, December 1903. In this paper, p. 417, the term “ wind tunnel” is used for the first time. ™ The resistance of the air determined at speeds below one thousand feet a second, with description of two new methods of measuring projectile velocities inside and outside the gun. Thesis, Johns Hopkins Univ., 46 pp., illus., 1808. “New methods of experimentation in aerodynamics; outline of some experi- ments made by H. Mattullath and A. F. Zahm, at the Catholic University of America. Paper communicated to the meeting of the American Association for the Advancement of Science, at Pittsburgh, June 30, 1902. 12 pp., illus., signed A. F. Zahm, Washington, D. C., 1902. NO. 4 PIONEER WIND TUNNELS—RANDERS-PEHRSON II epoch-making paper on ‘Atmospheric Friction”’,” read before the Philosophical Society of Washington, February 27, 1904. This paper disclosed for the first time the fact that skin friction is responsible for the major part of the total drag. The tests were made in the wind tunnel on carefully constructed boards up to 16 feet long suspended on the wire balance. Tests were also made on various spindle- and fish-shaped bodies, establishing the best form for airship hulls and giving, for the first time, the reason why the now universally accepted torpedo shape is preferable. The resistance of wires, struts, wings, and other airplane parts was also studied. The tunnel was also used for instruction at the University, several students taking part in the experiments. Occasionally, special tests were made for other investigators; for instance, Octave Chanute sent a stuffed buzzard for lift and drag measurements, and Emile Berliner had a monoplane model tested. WRIGHT BROTHERS The Wright Brothers’ gliding experiments at Kitty Hawk in 1901, although they seemed successful to other observers, were very dis- appointing to the Wrights themselves, as the new glider did not at all perform according to their calculations based on the aerodynamic tables of Lilienthal. On returning to Dayton in August, they decided to find out by laboratory methods what was wrong. Their first testing machine consisted of a bicycle wheel mounted horizontally on a spar projecting from the front of a bicycle. The relative aerodynamic efficiency of various surfaces was found by mounting them on this wheel, balancing one against the other and riding the bicycle at a fairly constant speed. Next they sent the blast from a fan through a square tube and mounted their surfaces as blades on a vane in the stream, balancing a curved surface against a plane surface. By the middle of October 1901 a small wind tunnel was completed. It was 16 inches square inside and about 6 feet long, with a glass top. The wind was forced through by a blower fan, and passed through a ** Atmospheric friction with special reference to aeronautics, pp. 237-276, diagrs., 1904. From Bull. Philos. Soc. Washington, vol. 14, 1904. Also printed in England: Atmospheric friction on even surfaces, with commentary note by the Rt. Hon. Lord Rayleigh, F. R. S. Reprinted from the Philos. Mag., July 1904, pp. 58-67, diagrs. IZ SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 honeycomb wind-straightener. The air speed was estimated to be 40 feet per second. The balance was based on the principle of using the normal pres- sure on a plane surface to measure the lift of an airfoil. The wing model and the normal plane were mounted on separate horizontal cross-stream bars so linked together that the wind lift on the model F1G.6.—Schematic drawing of the Wright Brothers’ balance, based on a photograph. tended to move it across stream. The drag on the normal plane would tend to resist this movement. When the two were exactly balanced, the ratio of lift to the resistance of the normal plane was indicated by a pointer. A balance of similar construction was later used by Orville Wright in his wind tunnel in Dayton.” * Warner and Norton, Wind tunnel balances. U. S. Nat. Adv. Comm. Aeronautics, Rep. no. 72, pp. 39-40, 1920. NO: 4: PIONEER WIND TUNNELS—RANDERS-PEHRSON 13 About 200 wing models made of sheet metal were tested in the wind tunnel. Each model was tested at 14 different angles of inci- dence, varying from 0° to 45°. Tests were also made to ascertain the effect of varying the aspect ratio, of superposing surfaces, etc. Great care was taken in making the tests; no one but the observer was allowed near the tunnel while it was in operation, and he kept the same position during the extent of the test, in order not to disturb the air current. The results were meticulously noted, and when completed they formed a valuable collection of aerodynamic tables which were later used by the Wrights as the basis for their design. Around Christmas 1902 these experiments came to an end, and the apparatus was taken down.” This construction and the Wright Brothers’ investigations there- with formed one of the chief factors leading to their success at Kitty Hawk on December 17, 1903. i. E. SPANTON The first wind tunnel at the National Physical Laboratory in Lon- don was set up several years before aeronautics became a subject of research at that institution. This predecessor of the great modern N. P. L. tunnels was built in 1903 by Dr. Thomas E. Stanton, for investigation of wind pressure on surfaces and structures. Stanton’s tunnel was vertical, the upper part a cylinder 2 feet in diameter and 44 feet long, terminating in a square box 4 by 4 feet, and 1 foot 3 inches deep where the balance was inserted. Underneath this, connected by a shorter length of pipe of the same diameter as the upper part of the tunnel, was the fan chamber. The fan, which produced the wind in the tunnel by suction, was driven by an electric motor and could be regulated to give air speeds from 5 to 30 feet per second. The balance comprising a horizontal lever carried on knife edges, had a sliding scale, and a scalepan with a dashpot for damping the vibrations. It was inserted in the center part of the tunnel, so that the model projected into the cylindrical section. The long arm of the lever was hollow and could be connected with a sensitive manometer * Aeronautical Journ., vol. 20, pp. 73-74, July-Sept. 1916. Unpublished letters of Wilbur Wright to Octave Chanute, now in the Library of Congress. l4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 for measurement of pressure. The speed of the current in various parts of the tunnel was measured by a pitot-static tube connected to the manometer. ELEVATION SECTIONAL PLAN Scace | IncH = 4 Feer FooT 1 ? ' z 3 4 7 Freer Fic. 7.—Stanton’s tunnel, 1903. Experiments were made in this very carefully constructed little tunnel to determine the resultant pressure and the distribution of pressure on round, square, and rectangular thin plates, normal and inclined to the current, on lattice work, cylinders, and finally on model roofs and bridges. The tunnel was still in use at the time when the 4-foot N. P. L. tunnel was constructed.” * Tnst. Civil Engineers, Minutes of Proc., vol. 156, pp. 78-139, illus., 1904. Great Britain Adv. Comm. Aeronautics. Rep. 1909-10, p. 14. NO. 4 PIONEER WIND TUNNELS—RANDERS-PEHRSON 15 GaAx. CROCCO In the fall of 1903 an aerodynamical laboratory was established near Rome by the “ Brigata Specialisti”’ of the 3rd Italian Engineer Corps. The laboratory was built under the direction of Lieutenant, now General, Gaetano Arthuro Crocco. It was well equipped with research apparatus, including, as the most prominent part, a wind tunnel of novel construction. By means of a 2.5-meter centrifugal fan and a 30-hp. electric motor, air was driven into a large cylindrical chamber like a gasometer, 5 meters in diameter and 3.5 meters high, which served to overcome turbulence and fluctuations. From here the wind was conveyed through a tunnel, the end of which was inserted in the laboratory wall, fan and air tank being outside. The cross-section of the mouth of the tunnel was I by I meter square; also a cross-section of 80 by 80 centimeters was used, and cylindrical mouthpieces of smaller diameter. The tests were made in the open jet, balance and models being mounted on a support on the floor, on a light carriage, or sometimes on floats swimming in water. The principal balance, constructed by Crocco in 1904, was an improvement on the dynamometric balance of Renard. For some researches the part of the laboratory where the air entered, was closed off to form a second air-straightening chamber, and the wind continued through a tunnel within the laboratory building. This tunnel was 8 meters long and 0.85 by 1.50 meters in cross-section.” The investigations at the laboratory of the Brigata Specialisti were chiefly concerning air propellers and the resistance and stability of airships. The construction of the first Italian military airship in 1907 was based on these tests. Results of the investigations were also presented in several im- portant papers by Crocco, published in various places, and reprinted in his “ Problemi aeronautici, degli albori fino alla guerra.” 524 p., illus., 27 pls. (Roma, A. Stock, 1931). D. RIABOUCHINSKY Through the efforts of D. Riabouchinsky, a wealthy patron of science and himself a scientist, Russia took its place in the front rank with regard to pioneer aerodynamic research. *” Boll. Soc. aeronautica ital., vol. 2, p. 209, illus,, Nov.-Dec. 1905. Marchis. Le Navire aerien. Appendix, pp. 122a-127a, illus. Paris, 1909. ioe) Oo’ J 2 > ‘C061 ‘JauuN} s,09I0IQV— 8s “OIY ban ee ee = = ee ee ee ' 200 BF 3) ies “eg y en ey ! oss? IS OS SSS SS SS SEA _—S—————__E————aaSSSSS} es aaa ra ae “7m Zo ei PUPP oP 217 Pr, 7 —— et ee eS —EEE srpssesz7 zucfas y 7972607 SUDp QUEUTZOT ' SMITHSONIAN MISCELLANEOUS COLLECTIONS 16 NO. 4 PIONEER WIND TUNNELS—RANDERS-PEHRSON 17 At Koutchino, not far from Moscow, Riabouchinsky erected, at his own expense, a complete aerodynamic laboratory with several buildings and an adequate staff. According to a suggestion by Prof. N. E. Joukovsky, the laboratory was equipped with a cylindrical wind tunnel, 14.50 meters long and 1.20 meters in diameter. Wind was produced by a suction fan, driven by an electric motor. Great pains were taken to render the wind uniform. After several experiments that did not give satisfactory results one end of the tunnel was en- closed in a cylindrical hood 2.2 meters in diameter and 3.5 meters long, coaxial with the tunnel itself. This admitted the air in such a way that a sufficiently even current was obtained. The testing section, in the middle of the tunnel, was provided with windows, from which the action of models could be observed. A great variety of aeronautical and hydrodynamical subjects were studied. Among the researches in the wind tunnel, the experiments with propellers, particularly lifting propellers were important. A bul- letin was published by the laboratory in six large issues, the last ii 1920... LUDWIG PRANDTL With the construction of the first wind tunnel at Gottingen we are approaching modern times. This was the first return-flow tunnel, built by Dr. Ludwig Prandtl for Motorluftschitfstudiengesellschaft and completed in July 1908." This tunnel was superseded in 1916-17 by a much larger tunnel with open jet and return flow, which is now called the Gottingen type. A. RATEAU With the aid of the Société d’Etudes de Locomotion Aérienne, A. Rateau built a wind tunnel in Paris in 1909. A 4-foot propeller blew the air into a rectangular chamber, 1.60 meters in cross-section, with an outlet contracting to a nozzle 70 centimeters square. The current was straightened by passing between a number of wooden ” Institut aérodynamique de Koutchino. 8 pp., 17 pls., St. Petersbourg, 1905. Institute aérodynamique de Koutchino, 1904-1914. Moscow, 1914. Bulletin de l'Institut aérodynamique de Koutchino. Fasc. 1-6. Moscow, 1909-1920. * Motorluftschiffstudiengesellschaft m. b. h. Berlin, Jahrbuch 1907/08-1912/13. Zeitschr. Ver. deutsch. Ingenieure, vol. 53, pp. 1711-1719, Oct. 16, 1909. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 slats, and the nozzle made possible speeds of up to 35 meters per second.” - GUSTAVE EIFFEL Gustave Eiffel built his first wind tunnel on the Champ de Mars in 1909. It was the open-jet nonreturn tunnel with an airtight testing chamber, known as the Eiffel type. The air current was cylindrical, I.5 meters in diameter, later enlarged to 2 meters.” In 1911 Eiffel moved to Auteuil and built a new and larger labora- tory, which he later turned over to the French Government. NATIONAL PHYSICAL LABORATORY The N. P. L. built its first large wind tunnel in London in 1910. It was 4 by 4 feet in cross-section and was supported inside another tun- nel 8 by 8 feet. The space between the walls of the two tunnels was a return passage for the air, which was drawn through the 4-foot tunnel by a Sirocco fan, driven by a 15 hp. engine.” This tunnel was not very satisfactory and was replaced in 1912 with a closed-jet nonreturn flow tunnel. The tunnels of Prandtl, Eiffel, and the N. P. L. have been very briefly described, as their main features are generally known, and full descriptions are readily available.” They end the pioneer period and begin a new era in wind-tunnel history. Before these three laboratories were established, powered flight had become a proved fact, and air- ships had met with considerable success. The necessity for reliable laboratory research soon became universally recognized and wind- tunnel laboratories were built and maintained by governments and institutions, as well as by private agencies. * Aérophile, vol. 27, pp. 266-268, illus., June 15, 1900. Soc. ingenieurs civils France, Mém. et C. R. Traveaux, vol. 65, pp. 61-78, illus., July 1912. ** Eiffel, Gustave, Installation d’un laboratoire d’aérodynamique. Paris, 1910. La résistance de l’air et l’aviation, experiences effectuées au Laboratoire du Champ de Mars. Paris, 1910. The resistance of the air and aviation, experi- ments conducted at the Champ de Mars Laboratory. Translated by Jerome C. Hunsaker. London and Boston, 1913. ** Great Britain Advisory Committee for Aeronautics, Rep. 1909-10, pp. 14-15, 2 folded plates. Flight, vol. 2, pp. 226-227, March 26, 1g1o. * Zahm, A. F., Report on European aeronautical laboratories, Smithsonian Misc. Coll., vol. 62, no. 3, 23 pp., 11 pls., 5 figs., 1914. 19 PIONEER WIND TUNNELS—RANDERS-PEHRSON NO. 4 UOEZIENSIA MOTT S]|!UupUur MA Siasuaspuod pue sjied suediry sdiysire JO poods [eo4a pue AqyIqe1s UOIINALAjSIP IINssotIg UOTeZI[eNsIA MOLT S[IOJILe UO S910 sodejins ourjd Jo Seip pue yy] yoreasat JO yoofqns JOJOU 911499]5] uey uoTjONS ‘dy Z 10}JOU! 914}99[H uey JoMolg ‘dy OO1 oulsua LWUea4S ue} JMO uey JaMOlg yeip Asuummyy SULSUd Ser) uey UOONS ainssoid ‘sq, OZ jof ureays SULSUD WRIIS ur} JaMO[q auIsUs pue tajjadwy sown], puryy Aaauorig fo suoynoyisads ‘Jas Jod tw o£'o “ay tod soy £ ‘Qas tod w Ol ty tod say zz ‘Jes Jad wi zz “ay tod sort 6 ‘Jas Jod wi FI “IY Jod soprum 1 ‘Qos tod wi SI “ry tod soprut pe ‘Jas Jod Ul O1 “ry tod sop ez ‘Das Jod UW QI “ay tod sopru 1 ‘Q9S Jod Ul QT “ay tod salut oF poeds sry uw OS"0 X 02 ‘Ul £61 XQ tw o£'0 x 0z'0 ‘UlLZI XQ “WIP WI 2 SO ‘WIRIp “Ul OF ® FOI ul 16'°0 x 10'0 Cg ooe (S W 0g0 “URI ‘UL Tf “WRI, ul £z'0 x S110 ‘ul6 x &V wi Sz’0 X QI'0 "ul ZO xZ uw v0 x $zo ‘ul ZI X OI Ul Qz"0 X 9z'0 ‘UI QI X QI UOT}OIS-SSOI) AdTeyy AdIE WW (sjouun} z) Inoje’] UX] ptreudy 130A pue 19SULUIT Oey sdryiyd weyua \\ otueN OOOI 6681 L081 g6gI 9681 POgI £6g1 VRgI IZgI a18q MISCELLANEOUS COLLECTIONS VOL. 93 SMITHSONIAN yoreasat otueuAposae [esauary yoreasat oueUu -Apoise [eioues pue sjlojiiy sioy[edoig ‘s[loyjiry yoieaset DWeUApOIe |etoues pue sjapow diysary yoreasat stueu -Apoloe [P1ouas pue SMoTOSITy ‘Oyo ‘sioy[adoig ‘sjepow diysiny sainjonijs uo dinssoid purA, ‘Oyo ‘orjes yoodse JO Poy “Seip pue FIT ‘Oyo ‘sq]nY ‘sjn44s *SSUIM UO Jd1OF Ae “UOIZOIAF UTYS Yoteasat JO Joo fqns ‘dy Sr 40}0W 914}99,q uey uoTons [esnyztsyuay ‘dy 99 10JOUI 914}99[H uey UOTONS ‘dy Sz 10jOUI 911499] ue} JOMOlT ‘dy PE 10}0W d14}D9[q uey uoTONS JOJOUL 911}99[ ue} UOTING JOJOW I11499[Y ue} JOMOTG [esNylayue) JOJOU 911399] uey UOT}INS ‘dy Z oulsua surposes) ue] JOMO[G ‘dy ZI JOJOW d4DaTyq uey UOTJINSG aUuISUa pue Jo[jadwy SJOUUN TT, puryy 4aau0ig fo suoynaifioads das Jod wi £1 “IY Jod sayrw o£ ‘Jas Jod wi 0z “IY tod sayru PP ‘Qas aod ur SE "ay dad say QZ ‘Qas Jod Wi OI “IY tod sapiw cz ‘Jas Jod wi oS'g “1y tod sapiut $V ‘Jas Jad wi 6z "1y tad sayrur $9 ‘Das tod wi 6 “IY tod solu oz ‘Jas Jod WzI “ay tod sai Zz ‘Jas Jod WII "IY tod saypiut Sz peeds Ivy WoT xe 1 yuxy Sa xeraiel ay cea ut 0Z°0 x 020 "ul $Zz x $Lz wZxe eS 9) “WIP WI ZI “‘WeIp Joop V wIXxi “ul O£ "WIRIp WW 19'0 ‘weIp ‘ul pe lw oO x OF'0 ‘UL OI X QI lw €g°1 x £Q'I Ce ete) UOT}I9S-SSO1I) “T'd 'N Jed ns neayey [pueig Aysuryonoqeny 099017 U0JURIS YS A\ WYPe7 alueN i 6001 6061 0001 go6r S061 £061 £061 1001 1001 97eq SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93, NO. 4, PL. 1 he 1. MAXIM’S TUNNEL, 1896 2. LA COUR’S TWO TUNNELS, 1897 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93, NO. 4, PL. 2 | MAREY’S TUNNEL, 1900 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93, NO. 4, PL. 3 1. A. F. ZAHM'S AERODYNAMIC LABORATORY, 1901 2. ZAHM'S TUNNEL WITH INTAKE CONE REMOVED SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93, NO. 4, PL. 4 Sh Nai ate ee RIABOUCHINSKY'S TUNNEL, 1905 ant ¥ _ Laie at a ae : 7 5 Tas, Pkt A j Hs an : 5 he Pi a yas | SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 5 NOMENCLATURE OF SOME CAMBRIAN TRILOBITES BY CHARLES ELMER RESSER Curator, Division of Invertebrate Paleontology, U.S. National Museum (PUBLICATION 3295) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION FEBRUARY 14, 1935 The Lord Baltimore Press BALTIMORE, MD., U. S&S A. NOMENCLATURE OF SOME CAMBRIAN TRILOBITES By CHARLES ELMER RESSER Curator, Division of Invertebrate Paleontology, U. S. National Museum INTRODUCTION After many years devoted to intensive study of Cambrian stratig- raphy and paleontology, the author hopes soon to submit for print- ing a complete summary of available information on the subject. Dur- ing the course of this research many interesting facts have been brought to light, and it becomes clear that much of the paleontology is not up to date. To include the necessary nomenclatural changes in the bibliographic portion of the summary would be to bury them; hence it is planned to publish, from time to time, separate papers embodying necessary changes. Dr. Charles D. Walcott’s intention to monograph the Cambrian trilobites as he did the brachiopods was repeatedly stated in his writ- ings and discussed in conversation over a period of many years. He finally came to realize that this would be more than a life-long work, since, while brachiopod species are numbered by hundreds, the trilo- bites comprise thousands. Unfortunately, just as he was well started on the description of the many new species in hand, matters pertain- ing to the World War robbed him of his research time. Although he accomplished much in this line, his assemblage of trilobites was so vast that many of the species remained unstudied and undescribed. With the study of the new material generic relationships formerly obscure become apparent, and a more satisfactory classification emerges. However, monographic studies of many more groups must be made before families or other groupings above generic rank can be attempted. For this reason the genera discussed in this paper are not referred to families. The data here presented do not lend themselves to precise syste- matic arrangement, for time is not available to properly monograph the genera or families discussed ; wherefore the information is given in a condensed form, and is arranged alphabetically by genera. How- ever, occasionally rather lengthy descriptions are presented, particu- larly when important generic questions are involved. Usually incom- SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 5 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 plete bibliographies are given, but they include references to all papers that add information. Care has been exercised to conform strictly to the rules of the International Commission on Zoological Nomen- clature. For the sake of lowering publication cost illustrations are omitted from this article, even though they would be desirable, par- ticularly since most papers describing Cambrian fossils are now out of print. New genera and species which require illustration to conform to the rules will be placed in a separate series of papers. Few foreign species are given consideration because several reports either in press or about to be printed care for many of them. ACROCEPHALITES Wallerius, 1895 Acrocephalites Wallerius, Unders zonen med Agnostus leavigatus i Vestergot- land, Sweden, p. 52, 1895. Acrocephalites Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 174, 1916. Much confusion exists respecting this genus, as many different trilobites have been referred to it simply because they possess a median boss. In fact, the genus was not understood until Wester- gaard * published photographs of the type species. From his studies it is evident that the North American forms cannot even belong to the same family, and consequently fall into other genera. Restricting the genus to species congeneric with the genotype, only three very rare forms remain, viz., Acrocephalites stenometopus (Angelin) and A. ? rarus Westergaard, from Sweden, and A. vigilans Walcott and Resser from Novaya Zemlya. Diagnosis—Cranidium alone known. Viewed vertically the facial suture converges forward from the posterior margin, but in side view it rises sharply from the basal plane of the head to the eyes and then drops off equally rapidly to the anterior angles. Cranidium keeled. Glabella large, about two-thirds as long as the head, tapering for- ward, but truncated anteriorly; two pairs of glabellar furrows turn sharply back; occipital furrow and ring well defined; sharp occipital spine. Brim with a well-defined rim about one-third the width of the preglabellar area; rim thickened and extended forward into a blunt spine medially. Fixed cheeks half as wide as the glabella; palpebral lobes strongly curved, elevated, and entirely beyond the rather straight course of the facial suture. Surface granulose. Genotype.—Solenopleura? stenometopa Angelin. Range.—The genus is confined to Upper Cambrian strata in the Atlantic Province. * Sveriges Geol. Unders., ser. Ca, no. 18, p. 123, pl. 1, figs. 20, 21, 1922. NO. 5 CAMBRIAN TRILOBITES—RESSER 3 Species formerly referred to Acrocephalites: A. americanus = Alokistocare A. insignis = Acrocephalops A. aoris = Alokistocare A. majus = Alokistocare A. aster = Deiracephalus A. multisegmentus = Deiracephalus A. glomeraius = Modocia A. tutus = cicrocephalops A. haynesi = Bolaspis A. vulcanus = Billingsaspis ACROCEPHALOPS Poulsen, 1927 Acrocephalops Poulsen, Meddels. Gr¢énland, vol. 70, p. 275, 1927. Numerous species formerly referred to Acrocephalites belong to this genus. At present all forms so identified are confined to the Ap- palachians and Greenland. Diagnosis —Cranidium with tapering glabella, furrowed; fixed cheeks wide, sometimes upturned from the well-defined dorsal fur- row ; eyes moderately small; eye lines heavy ; brim wide, with median boss (usually) and a narrow thickened rim of even width. Free cheeks with no peculiar characteristics. Surface granulated or lined. Thorax and pygidium not known. Comparisons.—Acrocephalops differs from Alokistocare primarily and chiefly in the presence of a thickened rim, narrow and of even width nearly to the anterior angles. From Bolaspis it is less clearly separated, the distinguishing criteria being the flatter shield, wider fixed cheeks, eyes apparently never stalked. Genotype.—A. gibber Poulsen. Range.—Possibly confined to the Middle Cambrian of the Appalachians and Greenland. DESCRIBED SPECIES REFERRED TO ACROCEPHALOPS Acrocephalops tutus (Walcott) Acrocephalites tutus Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 181, pl. 24, figs. 6, 6a, 1916. Middle Cambrian, Conasauga; (loc. 141) near Cave Spring, Georgia. Cotypes—U.S.N.M. nos. 61566, 61567. Acrocephalops insignis (Walcott) Acrocephalites insignis Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 179, pl. 25, fig. 1(only), 1916. (Not figs. 1a, 1b= A. nitida.) Again, several species were included under one name and must, therefore, be separated. The name insignis is retained for the forms with the strongest sculpturing. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Middle Cambrian, Conasauga; (loc. 112) 5 miles southeast of Center, Alabama. Lectotype and plesiotypes—U.S.N.M. no. 61568. Acrocephalops nitida, n. sp. Acrocephalites insignis Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 170, pl. 25, figs. 1a, 1b, 1916. (Not fig. 1= A. insignis.) Acrocephalites americanus Walcott, idem, pl. 24, fig. 3a. (This poor specimen may represent the species.) Unfortunately, none of the illustrated specimens is well preserved, and that shown in figure Ia has been damaged since it was photo- graphed. However, this form clearly has weaker eye lines and other sculpturing than A. insignis. Occurrence same as preceding. Cotypes—U.S.N.M. nos. 61569, 61570, and possibly 61560. ALOKISTOCARE Lorenz, 1906 Alokistocare Lorenz, Zeitschr. deutsch. Geol. Gesell., vol. 58, no. 1, p. 62, 1906. Alokistocare Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 182, 1916. Amecephalus Walcott, idem, vol. 75, no. 2, p. 53, 1924. Amecephalus Walcott, idem, vol. 75, no. 3, p. 65, 1925. If the rules of nomenclature did not require recognition of Alokis- tocare because Lorenz named a described species as its type, the original description would remain meaningless. Furthermore, the choice of A. subcoronatum as the genotype is unfortunate because only cranidia of the species are definitely known, and hence uncer- tainty prevails as to whether the tail was small or large, which in turn prevents final determination of generic limits. This problem is further complicated by the fact that cranidia similar to that of the genotype occur in entire specimens with both small and large pygidia. However, a search among the scores of cranidia of A. subcoronatum from Blacksmith Fork, Utah, reveals the presence of two pygidia which can represent the species, as they are small and similar to those of other species here referred to the genus. Much confusion exists between Alokistocare and Acrocephalites as well as among species of several other genera, because many strongly bossed trilobites which appear to belong rightly to Alokistocare have been assigned to the Atlantic Province genus Acrocephalites, and, as discussed above, their removal leaves no North American species in that genus. On the other hand, certain species must also be removed from Alokistocare to new genera. Owing to the confusion of species, Alokistocare has not been prop- erly understood, although in 1916 Walcott presented a diagnosis, evi- NO. 5 CAMBRIAN TRILOBITES—RESSER 5 dently based on other than the type species, which is correct in most essential points. Furthermore, it now appears that Amecephalus was not well founded and is congeneric with A. subcoronatum. The rela- tionship between A. subcoronatum and A. piochensis is apparent in the cranidium, and if the pygidium mentioned above represents J. subcoronatum, this part also conforms. In addition several entire in- dividuals belonging to undescribed species prove this relationship. A new generic diagnosis based on the enlarged group is given below. Diagnosis —Entire many-segmented trilobite tapering from a wide cephalon to a small pygidium. Cephalon semicircular, and usually with considerable convexity. Facial suture diverges only slightly in front of the eyes and is intra- marginal for perhaps one-third the distance to the center. Behind the eyes it diverges rapidly, forming short, blunt posterolateral limbs. Glabella marked off by well-defined dorsal furrow; tapers slightly and has three or four usually short glabellar furrows; length usually slightly more than half the length of the cranidium. Occipital furrow developed ; neck ring thickened. Brim wide; rim usually defined but sometimes by only a faint furrow; preglabellar area equal to or wider than the rim; and in common with many wide-brimmed trilo- bites usually has a more or less well developed median boss; brim striated vertically, anterior to the eye lines. Fixed cheeks wide, with large, strongly bowed palpebral lobes. Eye ridges usually strong ; eyes small, situated about the middle of the glabella. Free cheeks fairly large, concave toward the margin and with flat, sometimes concave genal spines. Thorax with approximately 20 segments, which are directed rather straight out, bending downward at the fulcrum and ending bluntly. Pygidium small, not exceeding in width the diameter of the glabella at the occipital furrow. Axis usually highly arched, as are also the pleural portions. Pleura fused; rim usually not demarcated. Surface striated vertically on the brim; some species are pustulose or granulose, sometimes with both a fine and a coarse set. Genotype.—Conocephalites subcoronatus Hall and Whitfield. Range——Chiefly in the Middle Cambrian, but also late Lower Cambrian. Species formerly referred to Alokistocare: A. labrosum = Bolaspis A. prospectense = Eldoradia A. linnarssoni = Eldoradia A, ticida = Bolaspis 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 DESCRIBED SPECIES REFERRED TO ALOKISTOCARE Alokistocare americanum (Walcott) Acrocephalites americanus Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 177, pl. 24, figs. 2, 2b, and possibly fig. 3, 1916. (Not 2a = A. georgense; 3a= Acrocephalops nitida?.) Two and perhaps three species are included in the original cotypes, and a brief examination of the nontype specimens from the locality indicates that perhaps twice that number of species occur. Conse- quently, the species must be carefully discriminated and restricted to the specimens conspecific with the lectotype. Middle Cambrian, Conasauga; (loc. 89) Livingston, Georgia. Lectotype-—U.S.N.M. no. 61557; paratype, no. 61559. Alokistocare georgense, n. sp. Acrocephalites americanus Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 177, pl. 24, fig. 2a, possibly also 3b, 1916. (See A. americanum.) Compared with A. americanum, this species has more granules, par- ticularly on the brim, and the rim is less thickened but upturned more sharply than shown in the illustrations. Occurrence same as preceding. Holotype —U.S.N.M. no. 61558. Alokistocare althea Walcott Alokistocare althea Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 184, pl. 25, figs. 3, 3a, 4, 5a, 1916. Middle Cambrian, Bright Angel: (loc. 74) Nankoweap Valley, and (loc. 74e) near Indian Garden Springs, Grand Canyon, Arizona. Cotypes—U.S.N.M. nos. 61571-61574. Alokistocare subcoronatum (Hall and Whitfield) Concephalites subcoronatus Hall and Whitfield, U. S. Geol. Surv. Expl. 4oth Parallel, vol. 4, p. 237, pl. 2, fig. 1, 18772 Ptychoparia subcoronata Walcott, U. S. Geol. Surv. Bull. 30, p. 205, pl. 28, fig. 4, 1886; idem, roth Ann. Rep., p. 652, pl. 96, fig. 6, 1891. Alokistocare subcoronatum Lorenz, Zeitschr. deutsch. Geol. Gesell., vol. 58, no. I, p. 62, fig., 1906. Alokistocare subcoronatum Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 187, pl. 25, fig. 2, 1916. Middle Cambrian, Ute; Blacksmith Fork Canyon, east of Hyrum, Utah. Cotypes.—U.S.N.M. no. 15442. Alokistocare aoris (Walcott) Acrocephalites aoris Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 178, pl. 26, figs. 3, 3b, 1916. NO. 5 CAMBRIAN TRILOBITES—RESSER Ve Middle Cambrian, Pleasant Hill; (loc. 107d) 1 mile northwest of Henrietta, Pennsylvania. Cotypes—U.S.N.M. nos. 61579, 61580. Alokistocare pomona Alokistocare pomona Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 186, pl. 25, fig. 6, 1916. It is not altogether certain that this species can remain in Alokis- tocare because of the narrow cranidium, but without additional speci- mens it cannot be determined how much of this is due to distortion of the soft shale matrix. Middle Cambrian, Park: (loc. 159f) Near Sixteen, Montana. Holotype-——U.S.N.M. no. 61577. Alokistocare majus (Walcott) Acrocephalites ? majus Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 180, Dl zowh eT TOLO: It appears that the median boss has been compressed laterally into a ridge. Middle Cambrian, Meagher; (loc. 4g) North of Gallatin River, east of Logan, Montana. Holotype-—U.S.N.M. no. 61578. Alokistocare piochense (Walcott) Ptychoparia piochensis Walcott (part), U. S. Geol. Surv. Bull. 30, p. 201, pl. 26, 2b; pl. 28, figs. 1, 1a, b, e, 1886. (Not pl. 26, 2a, nor pl. 28, Ic, 1d= A. packi: pl. 26, fig. 2= Glyphaspis nevadensis.) Ptychoparia piochensis Pack (part), Journ. Geol., vol. 14, p. 207, pl. 2, fig. 4a (only), 1906: (For other figures see below.) Liostracus piochensis Lorenz, Zeitschr. deutsch. Geol. Gesell., vol. 58, no. 1, p. 61, fig., 1906. Amecephalus piochensis Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 53, pl. 9, 1924. Amecephalus piochensis Walcott, idem, vol. 75, no. 3, p. 65, pl. 15, figs. 8-10, 1925. As originally set up this species was a mixture of specific and generic forms. As used by Walcott in 1886, plate 26, figure 2a, and plate 28, figures Ic and Id, represent another species, A. pack; plate 26, figure 2, represents Glyphaspis, which may be the same as that de- scribed by Pack under the name Ptychoparia kempi. Of Pack’s fig- ures, only 4a can possibly belong to Alokistocare; however, if this figure is correctly drawn, it is not piochensis, but an undescribed species. Figure 4 appears to be A. packi, 4b belongs to Glossopleura packi, and 4c to Clavaspidella howelli. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 At first glance this species, which was made the genotype of Amecephalus, is quite distinct from Alokistocare. Closer examination shows conclusively that it is congeneric with A. subcoronatum and must therefore be referred to it as the older genus. Because the speci- mens are considerably flattened, A. piochensis appears somewhat dif- ferent from A. subcoronatum, but if several specimens are carefully examined and variations due to preservation noted, generic distinc- tions are wanting. Description—Entire shield tapers from a wide cephalon to a small pygidium ; but because of the fewer thoracic segments, this feature is less pronounced than in other, longer, undescribed species. Cephalon semicircular in outline and apparently of moderate con- vexity. Cranidium about equally broad as long. Anterior facial su- ture diverges about 14°, to rather square anterior corners; intra- marginal possibly one-half the distance to the center; posterior por- tion diverges rapidly, forming long, narrow postero-lateral limbs. Glabella well defined by dorsal furrow, which, as usual, is shallower across the front and anterior to the eye lines ; tapers considerably and is a little over half as long as the cephalon. Three pairs of glabellar furrows traceable in some specimens. Occipital furrow shallow in the middle portion, separating a ring of even width. Brim wide, strongly striated vertically, anterior to the eye lines; a slight median boss usually discernible. When the free cheeks are retained, a narrow rim is apt to be present. In most individuals the doublure is pressed through so that its inner edge shows on the upper side as a ridge, with the consequence that many photographs seem to show a broad rim; doublure much wider in the center than toward the anterior angles, owing to the intramarginal course of the facial suture on the under surface. Fixed cheeks wide, eyes small, strongly bowed; eye ridges strong and, when undistorted, apparently fairly straight. Free cheeks also wide, with a concave border; sometimes with a narrow rim; genal spines concave and sharply pointed. Thorax with 19 segments. Pleura straight; fulcrum far out; fur- rows in a central position extending to the fulcrum. Pygidium small, with well-defined axis extending its full length. Pleura well fused, with pleural furrows showing only faintly on cer- tain moulds. Comparisons.—Confusion of this species with described forms is not likely because of its broad brim and widely placed fulcrum. Middle Cambrian, Chisholm ; (loc. 31) 3 miles northwest of Pioche, Nevada. Lectotype and paratypes —U.S.N.M. no. 15434b. _ NO. 5 CAMBRIAN TRILOBITES—RESSER 9 Alokistocare packi, n. sp. Ptychoparia piochensis Walcott, U. S. Geol. Surv. Bull. 30, p. 201, pl. 26, fig. 2a, pl. 28, 1c, 1d, 1886. (See A. piochensis.) In the narrowness of the cranidium this species is like A. pomona. Occurrence same as preceding. Holotype and paratype —U.S.N.M. no. go171. Alokistocare charax (Walcott) Ptychoparia charax Walcott, Smithsonian Misc. Coll., vol. 67, no. 2, p. 31, pl. 6, fie 1, LOZ: Ptychoparia pylas Walcott (part), idem. pl. 6, fig. 4b. This trilobite seems to belong to Alokistocare even though the fixed cheeks are somewhat narrow and eyes slightly large, in which respects it recalls Glyphasis, but since it fails to go far enough in that direction, reference to that genus does not seem to be warranted. In the pub- lished illustrations the eye lines are drawn too heavy. Middle Cambrian, Gordon; (loc. 4q, 4v) between Gordon and Youngs Creeks, Lewis and Clarke Range, Montana. Holotype —U.S.N.M. no. 63736; paratype, no. 63742. Alokistocare agnesensis (Walcott) Olenopsis ? agnesensis Walcott, Smithsonian Misc. Coll., vol. 57, no. 8, p. 242, pl. 36, fig. 2, 1912; idem (part), vol. 67, no. 3, p. 75, pl. 13, fig. 5, 1917. (Not 5a-c= A. stephenensis.) Middle Cambrian, Ptarmigan; (loc. 35m) 3 miles southwest of head of Lake Louise, Alberta. Holotype —vU.S.N.M. no. 58363. Alokistocare stephenensis, n. sp. Olenopsis ? agnesensis Walcott (part), Smithsonian Misc. Coll., vol. 67, no. 3, p. 75, pl. 13, figs. 5a-c, 1917. (Not 5, see preceding.) Middle Cambrian, Ptarmigan: (loc. 58k) north shoulder Mount Stephen, 3 miles east of Field, British Columbia. Alokistocare cleora (Walcott) Olenopsis cleora Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 74, pl. 13, fig. 3, 3a, 1917. Lower Cambrian, Mount Whyte; (loc. 62w) Gog Lake, Wonder Pass, British Columbia. Holotype-—U.S.N.M. no. 64396. Alokistocare stator (Walcott) Agraulos stator Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 173, pl. 36, fig. 6, 1916; idem, vol. 67, no. 2, p. 28, pl. 6, fig. 6, 1917. 1e) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 This species is not quite typical of the genus because the tips of the thoracic pleura are blunt, but this feature is hardly a generic criterion. Middle Cambrian, Ptarmigan; (loc. 35c) Mount Bosworth, British Columbia; (loc. 35m) 3 miles southwest of head of Lake Louise, Alberta. Holotype—U.S.N.M. no. 61729. ANORIA Walcott, 1924 Anoria Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 54, 1924; idem, no. 3, p. 67, 1925. Genotype.—Dolichometopus tontoensis Walcott. Range.—Middle Cambrian. DESCRIBED SPECIES REFERRED TO ANORTA Besides the genotype A. tontoensis, several other described species are now referred to the genus. Anoria bantius (Walcott) Bathyuriscus bantius Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 336, pl. 40, figs. 2-2c, 1916. Raymond suggested that this species might go into Athabascia, (=Clavaspidella), but it will be observed that, except for the exten- sion of the pleural furrows to the margin of the pygidium, it has no feature in common with Clavaspidella. Middle Cambrian, Rutledge; (loc. 12b) McAnnallys Ridge, 12 miles northeast of Knoxville, and (loc. 11) 1 mile east of Post Oak Spring, Tennessee. Cotypes—U.S.N.M. nos. 62661-4. Anoria bessus (Walcott) Dolichometopus ? bessus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 362, pl. 51, figs. 3-3c, 1916. Middle Cambrian, Park; (loc. 621) near Sixteen, Montana. Lectotype-—U.S.N.M. no. 62699; paratypes, nos. 62700-62701. Anoria baton (Walcott) Dolichometopus baton Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 362, pl. 51, figs. 2-2b, 1916. Middle Cambrian, Gordon; (loc. 3j) 6 miles northwest of Scape- goat Mountain, Powell County, Montana. Lectotype-—U.S.N.M. no. 62696; paratypes, nos. 62697-98. ee NO. 5 CAMBRIAN TRILOBITES—RESSER TE IL Anoria utahensis, n. sp. Bathyuriscus productus Walcott (part), U. S. Geol. Surv. Bull. 30, p. 217, pl. 30, figs. 1a, 1g, 1h, 1886. Sorting of the material from this locality identified as Bathyuriscus productus in 1886 and referred to B. anax in 1916, revealed the fact that neither mentioned species was present but that two other forms represented species referable to two other genera. Besides the illustrated head and pygidia, many other cranidia are ‘in hand as well as pygidia typical of Anoria. Middle Cambrian, Ophir; (loc. 30a) 1 mile below Argenta Cotton- wood Canyon, Wasatch Range, Utah. Holotype and paratypes—U.S.N.M. no. 15458. APHELASPIS, n. gen. A very abundant trilobite characterizing the upper beds of the Cap Mountain formation was described by Shumard as Conocephali- tes depressus. Recently, the genus has been found in beds of cor- responding age in Wisconsin and possibly in the western United States. Diagnosis —Cranidium rather long and narrow; rather flat. Facial suture diverges anterior to the eyes. Glabella narrow, slightly tapered ; glabellar furrows very faint; occipital furrow shallow but distinct. Brim wide; rim well defined, flat, usually somewhat upturned ; pre- glabellar area more than twice width of rim; usually rather convex. Fixed cheeks only about one-third width of glabella; eye lines clearly developed, eyes moderate in size, situated about center of head ; palpe- bral lobes strongly bowed, palpebral furrow distinct. Free cheeks rather large, long, with long genal spines; ocular platform swollen, elongate. Pygidium short, wide ; axis arched high above pleural lobes, extend- ing to rear margin; three axial furrows clearly defined ; pleural fur- rows less well developed. Genotype.—Conocephalites depressus Shumard. Name.—Agderes = neat ; aoms = shield. Range—Upper Cambrian. Above Crepicephalus zone and below Ironton. Aphelaspis depressa (Shumard) Conocephalites depressus Shumard, Amer. Journ. Sci. 2d ser., vol. 32, p. 219, 1861. Ptychoparia depressa Miller, North Amer. Geol. Pal., 1880, p. 565, (gen. ref.). I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Inasmuch as Shumard’s types were not illustrated and were later destroyed by fire, the propriety of recognizing his species may be questioned, but Walcott long ago set aside specimens as the species, and since they agree with Shumard’s description, little doubt of their identity remains. Upper Cambrian, Cap Mountain (Aphelaspis depressa zone) ; (loc. 67) Potatotop, 7 miles northwest of Burnet, Texas. Plesiotypes—U.S.N.M. no. 90172. ASAPHISCUS Meek, 1873 Asaphiscus Meek, 6th Ann. Rep. U. S. Geol. Surv. Terr., p. 485, 1873. Asaphiscus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 381, 1916. Asaphiscus was made in a footnote, without illustrations of the type species. Subsequently, the type and other species were well illus- trated, but as now constituted several distinct genera are included in Asaphiscus. Diagnosis.—Entire trilobite subelliptical. Cephalon semicircular in outline. Glabella wide, subconical, with only faint traces of glabellar furrows; occipital furrow also almost obsolete. Brim wide; rim wide, slightly thickened, flat, suture intramarginal for a short distance. Fixed cheeks narrow ; postero-lateral limbs triangular ; palpebral lobes moderately bowed. Free cheeks small, simple, without or with short genal spines. Nine thoracic segments in genotype. Pygidium nearly as large as cephalon. Pleural furrows weak ; when doublure shows, it appears to have a wide rim. Genotype—A. wheeleri Meek. Range.—Middle Cambrian. Species formerly referred to Asaphiscus: A.agatho = Genus undet. A, capella = Glyphaspis A.anaxis = Genus undet. A. duris = Genus undet. A. bradleyi = (new genus—Canadian A. florus = Genus undet. age) A. granulatus = W ecksina A. calanus = Coosia A. minor = Cedaria A. calenus = Glyphaspis A, unispinus = W eeksina A. camma = Glyphaspis DESCRIBED SPECIES REFERRED TO ASAPHISCUS At present Asaphiscus iddingsi Walcott (part) from Manchukuo is the only described species besides the genotype, remaining in the genus, but other new species are at hand. NO. 5 CAMBRIAN TRILOBITES—RESSER 13 Asaphiscus wheeleri Meek Asaphiscus wheeleri Meek, 6th Ann. Rep. U. S. Geol. Surv. Terr., p. 485, 1873. Asaphiscus wheeleri White, Rep. U. S. Geogr. Surv. West tooth Meridian, vol. 4, p. 43, pl. 2, figs. 1a-f, 1877. Asaphiscus wheeleri Walcott, U. S. Geol. Surv. Bull. 30, p. 220, pl. 31, figs. 3, 3a; pl. 25, fig. 9, 1886. Asaphiscus wheeleri Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 390, pl. 58, figs. I-1g, 1916. Meek did not figure this species when he described it. Later White figured six specimens, stating that they were those used by Meek. White’s figure 1f is the tail of an almost entire individual. The speci- men shown in figure Ie is lost, and it is possible that several other unmarked specimens were used by Meek. In 1886 Walcott presented two figures, the entire shield being a composite of Meek’s original cotypes. The head shown in figure 3a cannot be located. Walcott (1916) figured eight specimens. Figure I is an individual found by G. K. Gilbert and is unusually large and complete. Figure te is one of Meek’s original cotypes, whereas Ia is again a composite. The small head shown in Id is from locality toy near Marjum Pass, sev- eral miles from Antelope Springs. Unfortunately, this head was not prepared, and hence the figure is of little worth. All published figures show the rim as thickened, which it is not. Middle Cambrian, Wheeler; (loc. 4) Antelope Spring, House Range, Utah. Cotypes.—U.S.N.M. no. 8576 (6 specimens) ; plesiotypes, nos. 62754-62760. BATHYURISCUS Meek, 1873 Bathyuriscus Meek, 6th Ann. Rep. U. S. Geol. Surv. Terr., p. 484, 1873. Bathyuriscus Walcott, U. S. Geol. Surv. Bull. 30, p. 215, 1886. Bathyuriscus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 330, 1916. Bathyuriscus Raymond, Amer. Journ. Sci., 5th ser., vol. 15, p. 310, 1928. If the genus is restricted to the forms agreeing with the genotype, only a few described species remain, but many new species will be described from the more recent collections. As the genus is herein described, the species fall into two groups: those with two marginal spines at the anterior corners of the pygidium; and the typical group, which lacks them. Inasmuch as all other characters appear to agree, this feature is not regarded as of generic value. Genotype.—Bathyuriscus ? haydeni Meek Range.—Middle Cambrian. I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Species formerly referred to Bathyuriscus: B. anax = Clavaspidella, etc. B. lodensis = Glossopleura B. bantius = Anoria B. ornatus = Klotziella B. belesis = Glossopleura B. productus = Glossopleura B, belus = Clavaspidella B. parabola = Glossopleura B. bithus = Clavaspidella B. pupa= occidentalis = Poliella B. dawsoni= Kootenia B. rossensis = Ptarmingia B. howelli = Clavaspidella B. senectus = Bonnia Reed’s species from India, B. ? stolickai is not Bathyuriscus, and most, if not all, of the Chinese forms are in the same category. B. batis Walcott, which may come from Lower Cambrian strata has a crani- dium much too wide to remain in the genus, but the material is so poor that a new name is not now suggested. DESCRIBED SPECIES REFERRED TO BATHYURISCUS Besides the species discussed below, B. rotundatus (Rominger) be- longs to the typical group of the genus. TYPICAL, NONSPINED GROUP Bathyuriscus haydeni (Meek) Bathyurus ? haydeni Meek, 6th Ann. Rep. U. S. Geol. Surv. Terr., (1872) p. 482, 1873. Bathyuriscus haydeni Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 341, pl. 46, figs. 2-2b, 1916. Meek had very fragmentary material and but little has been found since, but the species is apparently correctly understood. Middle Cambrian, Meagher; north of the Gallatin River, east of Logan, Montana. Cotypes.—U.S.N.M. no. 7863. Bathyuriscus powersi Walcott Bathyuriscus (Poliella) powersi Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 351, pl. 46, fig. 1, 1916. Middle Cambrian, Meagher ; Pole Creek, Madison Range, Montana. Bathyuriscus obrutchevi (Lermontova) Proetus sculptus Korovin (not Barrande) 1924. (Reference not available.) Olenoides obrutchevi Lermontova, Bull. Com. Geol. Leningrad, vol. 44, no. 8, p. 764, pl. 18, figs. 1-10, 1926. Middle Cambrian ; Tcheremkhova, Irkutsk, Siberia. Bathyuriscus piedmontensis, n. sp. Bathyuriscus sp. undet. Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 348, pl. 49, fig. 3, 3a, 1916. ——— ee Oe eee NO. 5 CAMBRIAN TRILOBITES—RESSER 15 It is possible that the illustrated pygidium does not belong to the species. Middle Cambrian, Conasauga ; (loc. 16e) 1 mile southwest of Pied- mont, Alabama. Holotype—U.S.N.M. no 62665; paratype (°?), no. 62666. SPINED GROUP OF BATHYURISCUS It may prove desirable to separate these forms as a distinct genus. Bathyuriscus adaeus Walcott Bathyuriscus adaeus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 334, pl. 47, figs. 3-3c, 1916. In most respects this species is much like B. rotundatus, but the anterior pygidial segment is extended beyond the margin into spines. The published illustrations fail to show correctly the full length or direction of the spines, which turn backward and reach about to the posterior end of the axis. Further, this spine interrupts the rim and does not arise from it. The specimen from locality 61j identified as this form represents another species. Middle Cambrian, Stephen; (loc. 58j) Mount Stephen, near Field, British Columbia. Cotypes—U.S.N.M. nos. 62631-4. Bathyuriscus atossa Walcott Bathyuriscus atossa Walcott, Smithsonian Misc. Coll., vol. 64, no 5, p. 336, pl. 48, figs. 2-2b, 1916. Middle Cambrian, Spence; (loc. 55c) Liberty Canyon, west of Montpelier, Idaho. Lectotype-—U.S.N.M. no. 62642 ; paratypes, nos. 62643-4. Bathyuriscus marjumensis, n. sp. Bathyuriscus ? sp. undet. Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 348, pl. 65, fig. 5, 1916. Presumably this form falls within the spined group of Bathyuriscus. If it rightly belongs here, it is apparently the youngest species of the genus known. Middle Cambrian, Marjum; (loc. 11p) 24 miles southeast of Marjum Pass, House Range, Utah. Holotype —U.S.N.M. no. 62845. 4 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 BILLINGSASPIS, n. gen. Several Lower Cambrian species referred with a query to Acro- cephalites constitute a distinct generic group. Since the genotype is a species described by Billings, the hybrid name Billingsaspis seems appropriate. Diagnosis ——Cranidium on the whole truncate-conical in shape; facial suture converging slightly, but because the anterior angles are considerably depressed this feature is accentuated in dorsal view. Glabella tapers considerably ; in limestone specimens three pairs of re- flexed glabellar furrows traceable. Brim consists of a depressed pre- glabellar area and a raised and thickened rim ; some specimens have a small median boss. Eyes small, eye lines present. Fixed cheeks rather wide and rising from the dorsal furrow to the angular palpe- bral lobes. Comparisons.—Superficially, Billingsaspis resembles Bolaspis, ex- cept for the depression where the latter has the great boss in the pre- glabellar area. Further study is necessary to show whether this re- semblance indicates genetic relationship. Genotype.—Conocephalites vulcanus Billings. Range.—Possibly confined to Lower Cambrian. Billingsaspis vulcanus (Billings) Conocephalites vulcanus Billings, Rep. Geol. Vermont, vol. 2, p. 952, fig. 357, 1861 ; repeated Geol. Canada, p. 286, fig. 206, 1863; original republished, Pal. Fossils Canada, vol. 1, p. 14, fig. 17, 1865. Ptychoparia vulcanus Walcott, U. S. Geol. Surv. Bull. 30, p. 198, pl. 26, fig. 4 (not 4a), 1886 U. S. Geol. Surv. roth Ann. Rep., p. 653, pl. 96, fig. 4 (not 4a=“ Ptychoparia” miser?), 1891. Acrocephalites ? vulcanus Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 182, pl. 26, fig. 2, 1916. Lower Cambrian, Parker; (loc. 25) Parkers quarry, Georgia, and (loc. 26) Corman farm, east of Highgate Springs, Vermont. Cotypes—U.S.N.M. no. 15437. BLAINIA Walcott, 1916 Asaphiscus (Blainia) Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 393, 1916. This undoubtedly deserves full generic rank; in fact, it is doubtful whether it is even closely related to Asaphiscus. Genotype.—Asaphiscus (Blainia) gregarius Walcott. Range.—Middle Cambrian. Species formerly referred to Blainia: B. glabra = Blountia B. paula = Genus undet. NO. 5 CAMBRIAN TRILOBITES—RESSER 17 DESCRIBED SPECIES REFERRED TO BLAINIA Blainia gregaria (Walcott) Asaphiscus (Blainia) gregarius Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 304, pl. 62, figs. 1, 1b-17 (not 1a= B. centerensis), 1916. It is possible that, even with the specimen figured as ta eliminated because it has 10 instead of 9 segments, more than one species still remains. Middle Cambrian, Conasauga ; (loc. 90) 3 miles southeast of Center, Alabama. Lectotype and paratypes—U.S.N.M. nos. 62797 and 62791, 62793-6, 62798-62801. Blainia centerensis, n. sp. Asaphiscus (Blaimia) gregarius Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 394, pl. 62, fig. ra, I916. This form has Io instead of 9 thoracic segments. Occurrence same as preceding. Holotype —U.S.N.M. no. 62792. BOLASPIS, n. gen. A rather large group of Cambrian trilobites, usually referred to Acrocephalites and Alokistocare because of the highly developed boss in the center of the preglabellar field, has long needed a generic name. Diagnosis —Cranidium characterized by a conical glabella, with one or more pairs of rather sharply reflexed furrows; fixed cheeks wide and tumid; eyes small and sometimes stalked; large rounded or tri- angular elevation in preglabellar field; rim usually flat and erect, particularly in the middle; neck ring heavy, sometimes extended into a spine. Thorax and pygidium unknown. Comparisons —When Acrocephalites was first described from frag- mentary specimens, which fail to give a true concept of the genus, the conical glabella and the boss in the preglabellar area were the sole generic features apparent. However, after the genus came to be un- derstood, it was apparent that the American species have only a few superficial resemblances and are not related to it. Comparing Bolaspis with Alokistocare, we find little similarity ex- cept in the possession of the median boss. Bolaspis has a much more tapering glabella, its preglabellar area is reduced to triangular form by the down-turned anterior angles, and it lacks the vertical striations crossing the brim. Genotype—Alokistocare ? labrosum Walcott. Name.—Bodos= lump, and aoms= shield. Range.—Apparently confined to the Middle Cambrian. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 DESCRIBED SPECIES REFERRED TO BOLASPIS Bolaspis labrosa (Walcott) Alokistocare ? labrosum Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 184, pl. 25, fig. 5 only, 1916. (Not 5a=B. nethartensis.) Several hundred cranidia permit the selection of specimens showing all features. Description —Glabella conical, with deeply impressed occipital fur- row ; two and sometimes three pairs of sharply recurved glabellar fur- rows are distinguishable on certain individuals ; neck ring drawn out into a heavy and probably fairly long spine; brim diamond-shaped, separated from the fixed cheeks by rather heavy, deep furrows which pass forward and outward from the glabella; rim somewhat thick- ened, highly arched in middle and nearly erect ; frontal furrow strong ; preglabellar area triangular, rising into a high boss in the middle ; fixed cheeks tumid, rising considerably higher than the glabella, the abrupt slope into the preglabellar area at times serving as an eye ridge; eyes small, situated about the middle of the cranidium, and usually stalked. The surface is granulose ; the granules are fairly large and scattered, and are most numerous on the glabella. At present the free cheeks and pygidia are not located. Middle Cambrian, Meagher ; (loc. 5f) 11 miles south of Neihart. Montana. Lectotype and plesiotype —U.S.N.M. no. 61575. Bolaspis neihartensis, n. sp. Alokistocare ? labrosum Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 184, pl. 25, fig. 5a only, 1916. Among the cotypes of B. labrosa is a much shorter and more com- pact form, to which the name AB. neihartensis is given. Comparisons —Compared with B. labrosa, the brim is reduced in width, with the consequent loss of the prominent boss. Furthermore, the dorsal furrows are more shallow and the fixed cheeks less ele- vated, so that the whole cranidium is rather evenly convex in cross- section. Finally, the surface of the new species appears to be smooth. Occurrence same as preceding. Holotype and paratype —U.S.N.M. no. 61576. Bolaspis haynesi (Walcott) Acrocephalites haynesi Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 179, pl. 24, fig. 4b (upper figure only), 1916. (Not 4, 4a= B. errata.) Three species were illustrated as B. haynesi, which fact vitiates this as an example of a variable form, and it is necessary to restrict the NO. 5 CAMBRIAN TRILOBITES—RESSER 19 name to one of them. Furthermore, among the new collections there are at least four additional species. The upper of the figures on the slab shown in figure 4b agrees most closely with the description and becomes the lectotype of B. haynesi. This specimen evidently carried a large occipital spine and not simply a thickened neck ring as in the picture. Middle Cambrian, Meagher ; (loc. 20k) Pole Creek, Madison Range, Montana. Lectotype-—Cast U.S.N.M. no. 61503 (one specimen). (Original in Mus. Comp. Zool.) Bolaspis raymondi, n. sp. Acrocephalites haynesi Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 179, pl. 24, fig. 4b (lower specimen), 1916. This species differs from B. haynesi in being less granulose ; wider across the fixed cheek, particularly at the anterior angles; and in possessing a stronger boss. Occurrence same as B. haynest. Holotype-—Cast U.S.N.M. no. 61562. (Original in Mus. Comp. Zool. ) Bolaspis errata, n. sp. Acrocephalites haynesi Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 170, pl. 24, figs. 4, 4a, 1916. B. errata has almost completely lost the median boss, so that the con- sequently narrower preglabellar area gives it an aspect different from more typical forms of the genus. Occurrence same as B. haynesi. Holotype—Cast U.S.N.M. no. 61562. (Original in Mus. Comp. Zool.) Bolaspis ticida (Walcott) Alokistocare ticida Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 187, pl. 26, figs. 6, 6a, 1916. Until better preserved material becomes available, we can tenta- tively refer this species to Bolas pis. Middle Cambrian, Bloomington; (loc. 55s) Blacksmith Fork, 15 miles east of Hyrum, Utah. Cotypes.—U.S.N.M. nos. 61589, 61590. CEDARIA Walcott, 1924 Cedaria minor (Walcott) Asaphiscus minor Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 388, pl. 61, figs. 3-3b, 1916. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Upper Cambrian, Weeks; (loc. 300) Weeks Canyon, House Range, Utah. Lectotype and Paratypes—U.S.N.M. nos. 62778-80. CLAVASPIDELLA Poulsen, 1927 Clavaspidella Poulsen, Meddels. Grgnland, vol. 70, p. 277, 1927. Athabascia Raymond, Amer. Journ. Sci., 5th ser., vol. 15, no. 88, p. 311, 1928. This genus differs from the others of the group formerly referred to Bathyuriscus in having a more rapidly expanding glabella anterior to the eyes, and in its large tail in which the pleural furrows extend to the margin, increasing in width toward their extremities. Genotype.—C. sinupyga Poulsen. Range.—Middle Cambrian. Poulsen was uncertain as to the age of the Cape Frederick VII formation, but it seems clearly to represent a portion of the Stephen formation and is therefore Middle Cambrian. DESCRIBED SPECIES REFERRED TO CLAVASPIDELLA Poulsen described three species: Clavaspidella sinupyga Clavaspidella platyrrhina Clavaspidella quinquesulcata Raymond also referred to Athabascia: Athabascia ostheimeri Raymond Athabascia belus (Walcott) Athabascia glacialis Raymond Clavaspidella howelli (Walcott) Bathyuriscus howelli Walcott, U. S. Geol. Surv. Bull. 30, p. 216, pl. 30, figs. 2, 2a, 1886. Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 343, pl. 47, figs. 1-1b, 1916. Middle Cambrian, Chisholm: (loc. 31) 3 miles northwest of Pioche, Nevada. Lectotype and paratypes —U.S.N.M. no. 15457. Clavaspidella bithus (Walcott) Bathyuriscus ? bithus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 340, pl. 47, figs. 4, 4a, 1916. Unfortunately, only pygidia of this species have yet been located. When a study of the large quantity of material from this locality is undertaken, heads will likely be found. Middle Cambrian, Spence; (loc. 55c) Liberty Canyon, West of Montpelier, Idaho. Lectotype-—U.S.N.M. no. 62635 ; paratype, no. 62636. NO. 5 CAMBRIAN TRILOBITES—-RESSER 2 Clavaspidella anax (Walcott) Bathyuriscus anax Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 335, pl. 48, figs. I, ta, 1c, Id (not 1b), 1916. A mere glance at the illustrations shows that the forms from Cotton- wood Canyon referred to this species belong to Glossopleura. Middle Cambrian, Ophir; (loc. 55e) Wasatch Range, north of Brigham City, Utah. Lectotype —U.S.N.M. no. 62637; paratypes, nos. 62638-40. Clavaspidella sylla (Walcott) Bathyuriscus (Poliella) sylla Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 354, pl. 65, figs. 2, 2a, 1916. This species approaches the extreme limits of the genus, but until more forms become available it seems inexpedient to make another genus. Middle Cambrian, Marjum; (loc. 110) 4 miles southeast of Ante- lope Springs, House Range, Utah. Cotypes—U.S.N.M. nos. 62837-8. COOSIA Walcott, 1911 Coosia calanus (Walcott) Asaphiscus calanus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 384, pl. 61, figs. 8, 8a, 1916. Upper Cambrian, Nolichucky; (loc. 47h) Wolf Creek, Bland County, Virginia. Cotypes—U.S.N.M. nos. 62789-90. DEIRACEPHALUS, n. gen. Several Upper Cambrian forms previously referred to Acrocephali- tes may be descendants of Middle Cambrian forms but are generi- cally distinct. Diagnosis —Cranidium quadrate in general outline. Sutures diverg- ing slightly anteriorly. Glabella tapering, rather conical, somewhat over half as long as the cranidium; no glabellar furrows visible on available specimens. Brim wide with a narrow rim of even width; preglabellar area crossed by a vertical median ridge, beginning at the dorsal furrow and widening out to join the brim. Eye lines strong ; palpebral lobes small, situated a little behind the middle of the cra- nidium. Occipital furrow separating the neck ring, which bears a spine. A beautiful compound eye has been preserved on a free cheek. Sur- face richly ornamented by pustules, or pustules and lines. 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL, 93 Comparisons —Compared with the somewhat older species of Alokistocare and Acrocephalops, this genus resembles both in certain features. Features of the rim as well as the median ridge, which re- places the boss, distinguish it from Acrocephalops, and the less ex- panded brim and the well-defined rim separate it from Alokistocare. Genotype—Acrocephalites ? aster Walcott. Name.—Aewpas = ridged, xehados = head. Range.—Possibly confined to the Upper Cambrian Crepicephalus zone. DESCRIBED SPECIES REFERRED TO DEIRACEPHALUS Deiracephalus aster (Walcott) Acrocephalites ? aster Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 178, pl. 26, figs. ob, oc, 1916. (Not figs. 9, 9a = L. buttsi.) Upper Cambrian, Conasauga; (loc. 22y) opposite car barn, Birm- ingham, Alabama. Holotype —U.S.N.M. no. 61594. Deiracephalus buttsi, n. sp. Acrocephalites ? aster Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 178, pl. 26, fig. 9, 9a, 1916. This form is similar to L. aster but differs in having fewer pustules and a heavier occipital spine. Upper Cambrian, Nolichucky ; (loc. 107¢) west base Copper Ridge, Tr miles northwest of Knoxville, Tennessee. | Holotype —U.S.N.M. no. 61593. Deiracephalus multisegmentus (Walcott) Acrocephalites multisegmentus Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 180, pl. 24, figs. 5, 5a, 1916. Upper Cambrian, Weeks; (loc. 30n or 300) 2 miles south of Marjum Pass, House Range, Utah. Cotypes—U.S.N.M. nos. 61564-5. DOLICHOMETOPUS Angelin, 1852 Dolichometopus Angelin, Pal. Scandinavica, pt. 1, Crustacea formationis transi- tionis Lipsiae, p. 72, 1852. Idem, Pal. Scandinavica, 3d ed. Holmiae, p. 72, 1878. Dolichometopus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 355, 1916. Numerous trilobites belonging to many genera have been referred to Dolichometopus, Corynexochus, and Bathyuriscus simply because their glabellae tended toward a rectangular form and expand forward. Angelin referred the first two genera to his family Corynexochidae. itt, deta step a a ~ e NO. 5 CAMBRIAN TRILOBITES—RESSER 23 No doubt the family is valid, but it must be confined to Dolichometopus and Corynexochus, which occur only in the Atlantic Province ; hence all other American and Asiatic forms must go into other genera. Properly restricted, Dolichometopus contains the species D. acadicus Matthew and D. suecicus, the genotype. The following list shows the generic position of the North Ameri- can forms not belonging to the Atlantic Province : D. baton = Anoria D. lodensis = Glossopleura D. bessus = Anoria D. occidentalis = Poliella D. bion = Glossopleura D. productus= Glossopleura D. boccar = Glossopleura D. tontoensis = Anoria D. expansus = (Not determined) D. varro = Housia The Oriental species ascribed to the genus all belong elsewhere. Their revision is now under way. DUNDERBERGIA Walcott, 1924 Dunderbergia Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 56, 1924; idem, no. 3, p. 85, 1925. Several species from Nevada, other than the genotype, D. mitida, appear to belong to the genus. Undescribed species from other regions also appear to belong to Dunderbergia. Genotype.—Crepicephalus (Loganellus) nitidus Hall and Whit- field. Range.—Upper Cambrian beds representing the Pterocephalia zone. DESCRIBED SPECIES REFERRED TO DUNDERBERGIA Dunderbergia maculosa (Hall and Whitfield) Crepicephalus (Loganellus) maculosus Hall and Whitfield (part), U. S. Geol. Expl. goth Parallel, vol. 4, p. 215, pl. 2, figs. 24, 25?, 1877. (Not fig. 26 = D, halli.) Ptychoparia maculosus Walcott, U. S. Geol. Surv. Monogr. 8, p. 269, 1884. Upper Cambrian, Secret Canyon ; (loc. 62a) opposite Jackson Mine, Eureka District, Nevada. Holotype-—U.S.N.M. no. 90670. Dunderbergia halli, n. sp. Crepicephalus (Loganellus) maculosus Hall and Whitfield (part), U. S. Geol. Expl., goth Parallel, vol. 4, p. 215, pl. 2, fig. 26, 1877. (See D. maculosa.) This figured pygidium is not from the same locality as the head of D. maculosa, and further, since the cranidia associated with this tail are also distinct, the logical procedure is to combine the two as a new species. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Upper Cambrian, Secret Canyon; (loc. 60) near Richmond Mine, Eureka District, Nevada. Holotype and paratypes —U.S.N.M. no. 24617. Dunderbergia simulator (Hall and Whitfield) Crepicephalus (Loganellus) simulator Hall and Whitfield, U. S. Geol. Expl. 4oth Parallel, vol. 4, p. 218, pl. 2, figs. 16-18, 1877. Inouyia simulator Walcott, Smithsonian Misc. Coll., vol. 64, 1916. Upper Cambrian, Secret Canyon ; Eureka District, Nevada. Lectotype-——U.S.N.M. no. 24575; paratypes, no. 24573. Dunderbergia granulosa (Hall and Whitfield) Crepicephalus (Loganellus) granulosa Hall and Whitfield, U. S. Geol. Expl. 4oth Parallel, vol. 4, p. 214; pl. 2, figs. 2, 3, 1877. Ptychoparia granulosa Walcott, U. S. Geol. Surv. Monogr. 8, p. 57, 1884. Inouyia granulosa Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 204, 1916. Upper Cambrian, Secret Canyon; (loc. 61) near Hamburg Mine and other localities, Eureka District, Nevada. Holotype —U.S.N.M. no. 24573. Dunderbergia pustulosa (Hall and Whitfield) Ptychaspis pustulosa Hall and Whitfield, U. S. Geol. Expl. goth Parallel, vol. 4, D223) pleen ties 27518775 Upper Cambrian, Secret Canyon; Pogonip Mountain, White Pine District, Nevada. Holotype-—vU.S.N.M. no. 24579. Dunderbergia suada (Walcott) Ptychoparia suada Walcott, Proc. U. S. Nat. Mus., vol. 13, p. 274, pl. 21, fig. 9, 1890. Upper Cambrian, Wilberns; (loc. 70) Baldy Mountain, near Mor- gans Creek, 8 miles northwest of Burnet, Texas. Holotype -—U.S.N.M. no. 23860. EHMANTIA, n. gen. Many Middle Cambrian species characterized by a peculiar pygidium have been referred to various genera during the past 55 years. When they are gathered together and their features noted, a definite, clear-cut genus emerges. To this is assigned the name of Philip Ehman, of Logan, Montana, who has been of material assistance in restudying the classic region along the north side of the Gallatin Valley. Diagnosis —Cranidium of a very common type. Glabella tapered, rounded in form, distinctly demarcated by dorsal furrow; glabellar NO. 5 CAMBRIAN TRILOBITES—RESSER 25 furrows usually very faint. Brim variable in width, with a convex pre- glabellar area and a flat, upturned rim. Eyes moderate in size, not much bowed, situated about the middle of the cranidium. Fixed cheeks about half as wide as the glabella. Free cheeks show suture intra- marginal for some distance and have a stout, short genal spine. Thorax has 12 to 14 segments in the specimens observed. Pygidium wide ; axis well defined except at rear; up to six or more axial rings are marked out. Pleural lobes very distinctive because both the pleural furrows and grooves are distinctly impressed to the very margin. Genotype.—Ehmania weedi, n. sp. Range.—Middle Cambrian. DESCRIBED SPECIES REFERRED TO EHMANIA Ehmania gallatinensis (Meek) Conocoryphe (Ptychoparia) gallatinensis Meek, 6th Ann. Rep. U. S. Geol. Surv. Terr., p. 485, 1873. Unfortunately this species is not illustrated, and it is necessary to restrict it to one of the three or more species present in the type lot. Middle Cambrian, Meagher ; north of Gallatin River, near Logan, Montana. Holotype —U.S.N.M. no. 7862. Ehmania walcotti, n. sp. Ptychoparia antiquata Walcott (part) (not Salter), U. S. Geol. Surv. Monogr. 32, pt. 2, p. 456, pl. 65, fig. 7a, 1899. (Not fig. 7= E. weedi.) This species was described as a variety. Many fine cranidia and pygidia occur among the specimens not illustrated. Middle Cambrian, Meagher; (loc. 151d) south of Gallatin River, Crowfoot Ridge, Yellowstone National Park, Wyoming. Cotypes—U.S.N.M. no. 90667. Ehmania weedi, n. sp. Genotype Ptychoparia antiquata Walcott (part) (not Salter), U. S. Geol. Surv. Monogr. 32, pt. 2, p. 456, pl. 65, fig. 7 (not 7a), 1899. This species has more strongly developed glabellar furrows than the other described species. Occurrence same as preceding. Holotype —U.S.N.M. no. 35234. Ehmania oweni (Walcott) Ptychoparia oweni Walcott (not Meek and Hayden), U. S. Geol. Surv. Mongr. 8, p. 55, pl. 10, figs. 3, 3a, 1884. (Probably also fig. 18, which is placed with Eteraspis laeviceps; and fig. 22.) 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Middle Cambrian, Eldorado ; east side Secret Canyon, Eureka dis- trict, Nevada. Cotypes—U.S.N.M. no. 24610. Ehmania smithi (Walcott) Anomocarella smithi Walcott, Smithsonian Misc. Coll., vol. 57, no. 4, p. 92, pl. 17, figs. 3-3a, 1911. Research in China, vol. 3, Carnegie Inst. Publ. 54, p. 203, pl. 19, figs. 16-16), 1913. Middle Cambrian, Conasauga; (loc. 90x) 3 miles east of Center, Alabama. Cotypes—U.S.N.M. nos. 58296, 582098. ELDORADIA, n. gen. Still another group formerly referred to Alokistocare deserves a separate generic name, and since it seems to reach its best develop- ment in the Eldorado limestone of the Eureka District, Nevada, may appropriately be named for the formation. Diagnosis.—Cranidium generally rectangular in outline, with a tapering glabella about half as long as the cranidium; usually well defined by shallow dorsal furrows; glabellar furrows faint or absent. Brim wide, usually with a large median boss; rim not sharply sepa- rated owing to the poor development of the anterior furrow. Fixed cheeks usually wide, with small, sometimes stalked eyes about op- posite the anterior end of the glabella. Comparisons—Compared with Bolaspis and Acrocephalops, this genus lies between the two but nearer to the former. From this it differs in greater width of fixed cheek, less clearly defined rim and shallower furrows. Genotype.—Ptychoparia ? linnarssoni Walcott. Range.—Presumably confined to the Middle Cambrian. DESCRIBED SPECIES REFERRED TO ELDORADIA Eldoradia linnarssoni (Walcott) Ptychoparia ? linnarssoni Walcott (part), U. S. Geol. Surv. Monogr. 8, p. 47, pl. 9, fig. 18a, 1884. (Not 1%=E. lata.) Alokistocare linnarssoni Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 185, pl. 25, fig. 7, and possibly the cheek inset in fig. 7a, 1916. (Not 7a=E. lata.) The best cranidium was not figured. Middle Cambrian Eldorado; (loc. 58) east side of New York Canyon, Eureka District, Nevada. Cotypes—vU.S.N.M. no. 24611. NO. 5 CAMBRIAN TRILOBITES——RESSER 27 Eldoradia lata, n. sp. Ptychoparia ? linnarssoni Walcott (part), U. S. Geol. Surv. Monogr. 8, p. 47, pl. 9, fig. 18, 1884. Alokistocare linnarssoni Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 185, pl. 25, fig. 7a (heads only), 1916. This species differs from EF. linnarssoni in its wider fixed cheeks. Occurrence same as E. linnarssoni (loc. 58a). Cotypes—U.S.N.M. no. 90669. Eldoradia prospectensis (Walcott) Ptychoparia ? prospectensis Walcott, U. S. Geol. Surv. Monogr. 8, p. 46, pl. 9, fig. 20, 1884; U. S. Geol. Surv. Bull. 30, p. 202, pl. 27, fig. 5, 1886. Alokistocare prospectense Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 186, pl. 25, fig. 8, 1916. Middle Cambrian, Eldorado; (loc. 52a) Prospect Mountain, Eu- reka District, Nevada. Holotype —U.S.N.M. no. 15441. ELRATHIA Walcott, 1924 Elrathia Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 56, 1924; idem, no. 3, p. 87, 1925. Already species have been referred to Elrathia improperly, as they formerly were to Ptychoparia, consequently great care must be ex- ercised to prevent this name also becoming a “ dumping ground.” These forms have what might be called the “ median” structure of the trilobite, consequently many genera are close to one another. Genotype.—Conocorphe (Conocephalites) kingi Meek. Range.—Likely confined to the Middle Cambrian. ADDITIONAL SPECIES REFERRED TO ELRATHIA Elrathia candace (Walcott) Ptychoparia candace Walcott, Smithsonian Misc. Coll., vol. 67, no. 2, p. 28, ple 6, figs: 35/30, 1017. Middle Cambrian, Gordon ; (loc. 4v) Gordon Creek, Powell County, Montana. Cotypes——U.S.N.M. nos. 63738, 63739. Elrathia pylas (Walcott) Ptychoparia pylas Walcott (part), Smithsonian Misc. Coll., vol. 67, no. 2, p. 33, pl. 6, figs. 4, 4a, 1917. (Not 4b= Alokistocare charax; 4c=— A. gordonensis. ) Middle Cambrian, Gordon; (loc. 4q) between Gordon and Youngs Creeks, Powell County, Montana. Cotypes—U.S.N.M. nos. 63740, 63741. 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Elrathia haguei (Hall and Whitfield) Crepicephalus (Loganellus) haguei Hall and Whitfield, U. S. Geol. Expl. 40th Parallel, vol. 4, p. 210, pl. 2, figs. 14, 15, 1877. Ptychoparia haguei Walcott, U. S. Geol. Surv. Bull. ro, p. 36, pl. 6, fig. 6, 1884. Species must be confined to the locality given below. Middle Cambrian, Eldorado; Pogonip Mountain, White Pine Dis- trict, Nevada. Holotype-—U.S.N.M. no. 24660. EUREKIA Walcott, 1924 Eurekia Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 56, 1924; idem, no. 3, Pp. 90, 1925. Genotype.—E. granulosa Walcott. Range.—Late Upper Cambrian. Previously the genus contained five species : Eurekia granulosa Walcott Eurekia eos (Hall) Eurekia angustifrons (Walcott) Eurekia binodosa (Hall) Eurekia dissimilis (Walcott) ADDITIONAL SPECIES REFERRED TO EUREKIA Eurekia denticulata (Meek) Proetus (Phaeton) denticulatus Meek, Rep. U. S. Geol. Expl. 40th Parallel, vol. 4, pt. I, p. 49, pl. 1, fig. 10, 1877. Originally assigned to the Devonian. Upper Cambrian, Hamburg; drift west side of Steptoe Valley, Nevada (float). Holotype —U.S.N.M. no. 14579. Eurekia finkelnburgi (Clark) Bayfeldia finkelnburgi Clark, Bull. Amer. Pal., vol. 10, no. 41, p. 32, pl. 4, fig. 7, 1924. Upper Cambrian, Norwalk sandstone ; Osceola Mills, Wisconsin. Holotype.—M.C.Z. no. 1712. ETERASPIS, n. gen. Diagnosis —Cranidium strongly arched longitudinally. Glabella poorly defined ; without furrows. Neck ring thickened ; occipital fur- row very shallow. Brim about one-third length of glabella; wide rim separated by shallow anterior furrow. Eyes moderate in size and moderately bowed, situated about middle of cranidium. NO. 5 CAMBRIAN TRILOBITES—RESSER 29 Pygidium flat. Axis defined by shallow dorsal furrow, but chiefly by arching above pleural lobes; extends to border. Pleural furrows faintly outlined. Border flattened. Genotype.—Ptychoparia laeviceps Walcott. Range.—Middle Cambrian. Name.—Erepos = another ; aoms=shield. Eteraspis laeviceps (Walcott) Ptychoparia laeviceps Walcott, U. S. Geol. Surv. Monogr. 8, p. 54, pl. 10, fig. 17, 1884. (Not 18= Ehmania oweni?.) Middle Cambrian, Eldorado; (loc. 58) east side Secret Canyon, Eureka District, Nevada. Holotype and paratypes —U.S.N.M. no. 24614. GLOSSOPLEURA Poulsen, 1927 Glossopleura Poulsen, Meddels. Gr¢gnland, vol. 70, p. 268, 1927. Diagnosis —Cephalon and pygidium large, of about equal size. Glabella wide, extending to anterior margin, where a narrow upturned brim may sometimes be present; glabella sometimes expands some- what forward of the eyes; four faint pairs of glabellar furrows usually traceable. Fixed cheeks small; eyes large, situated well back. Free cheeks rather wide with a broad rim when the wide doublure is im- pressed on them, otherwise sloping smoothly to margin. Thorax has seven segments. Pygidium usually has the axis well defined. Axial furrows often faint; pleural furrows faint but clearly traceable in some species to outer edge ; wide rim sometimes definite, when doublure is impressed on test. Genotype.—Dolichometopus boccar Walcott. Comparisons.—Glossopleura, as pointed out by Poulsen, resembles Anoria in the number of thoracic segments and the extension of the glabella to the front margin, but differs in having longer eyes. How- ever, his observation that Glossopleura lacks the macropleural de- velopment of the fifth thoracic segment does not hold if the present specific references are all correct. From the other members of the family Glossopleura it is distinguish- able by its less developed furrows, the absence of a brim on the cephalon, and the large size and posterior position of the eyes. Range.—Apparently confined to the Middle Cambrian of Green- land, the Appalachians, and the Rocky Mountains. Frequently the species of this genus constitute the entire fauna, except that an Alokistocare may go with it. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 DESCRIBED SPECIES REFERRED TO GLOSSOPLEURA The four species from Greenland described by Poulsen appear to be typical of the genus. ; Glossopleura boccar (Walcott) Dolichometopus boccar Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 363, pl. 52, fig. 1a, Ic, Ie, 1916. (Not fig. 1=G. stephenensis; fig. 1b=G. bosworthensis; figs. 1d, 1f = G. nitida.) Glossopleura boccar Poulsen, Meddels. Grgnland, vol. 70, p. 268 (gen. ref.) 1927. It appears that four species are represented among the figured specimens of D. boccar, consequently one form must be chosen as the species, in order that the genotype of Glossopleura can be understood. Keeping in mind Walcott’s habit of always writing the type locality first, and the statement he makes on page 363 that D. boccar is from the Mount Bosworth section, we may at once conclude that the latter is the type locality and hence choose the types of D. boccar from the illustrated specimens of locality 57g. This leads us to use figures 1a, Ic, Ie, as the cotypes of G. boccar. Figure ta shows the cranidial features imperfectly, first, because in the specimen the frontal ex- tension of the glabella is bent down by rock folding, and second, the photograph was cut off too short. Examination of the specimen, how- ever, reveals that its glabella is essentially like that of Figure tc. The specimens not illustrated from other localities referred to the species with more or less reservation represent new forms. Numerous unworked collections have been made from this zone, and it is pos- sible that the species may be found in them, but a cursory study indi- cates that most belong to new forms. Description—Head and tail of even size, semicircular in outline. Glabella nearly rectangular, but slightly constricted behind the center ; glabellar furrows faint. Fixed cheeks practically confined to the palpe- bral lobes ; none in front of the eyes. Eyes long and situated far back. Thorax with seven segments. Pygidial axis rather high and well marked by the dorsal furrow. Five or six axial furrows distinguishable. Doublure wide, and when pressed against test, makes a well-defined border; otherwise the pleural lobes slope rather steeply down to the margin all around. Pleural furrows traceable but not deep. Middle Cambrian, Stephen; (loc. 57g) Mount Bosworth, British Columbia. Cotypes—U.S.N.M. nos. 62703, 62705, 62707. —_ = NO. 5 CAMBRIAN TRILOBITES—RESSER 31 Glossopleura bosworthensis, n. sp. Dolichometopus boccar Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 363, fig. 1b only, 1916. (See preceding, G. boccar.) Several cranidia and a hypostoma appear to constitute another species. Compared with G. boccar, the glabella is even more nearly rectangular, the eyes slightly longer and directed more outward. Occurrence same as G. boccar. Holotype-—U.S.N.M. no. 62704. Glossopleura nitida, n. sp. Dolichometopus boccar Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 363, pl. 52, figs. 1d and 1f only, 1916. (See preceding, G. boccar.) Cranidia of this species are much shorter than the others formerly referred to G. boccar. The hypostoma is placed with the illustrated pygidium because its surface ornamentation is similar. Occurrence same as G. boccar. Holotype—U.S.N.M. no. 62708; paratype, no. 62706. Glossopleura stephenensis, n. sp. Dolichometopus boccar Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 363, pl. 52, fig. 1 only, 1916. (See preceding, G. boccar.) Only this one impression representing the genus has been found among the thousands of specimens collected from the Mount Stephen fossil bed. Unfortunately, it is not well preserved. Comparisons —Compared with D. boccar, to which it had been re- ferred, this species has a shorter glabella without glabellar furrows, and the dorsal furrow is more curved, causing greater expansion of the glabella at both ends. In the tail axial and pleural furrows are practically absent. Middle Cambrian, Stephen; (loc. 14s) Mount Stephen, near Field, British Columbia. Holotype—U.S.N.M. no. 62702. Glossopleura producta (Hall and Whitfield) Ogyia producta Hall and Whitfield, U. S. Geol. Expl. goth Parallel, vol. 4, p. 244, pl. 2, figs. 31-34, 1877. Bathyuriscus productus Walcott (part), U. S. Geol. Surv. Bull. 30, p. 217, pl. 30, fig. 1f and composite from two localities, 11, 1886. (Not Ic, 1d=G. piochensis; 1=G. parabola; ta, 1g, th=Anoria utahensis ; 1b=G. utahensis.) Dolichometopus productus Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 369, pl. 53, figs. 2-2e only (all copies of originals), 1916. Many species have been referred to this form. Every occur- rence except unquestioned Ophir shale can be eliminated without examination, 3 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Middle Cambrian, Ophir, East Canyon, and (loc. 3c) Ophir, Oquirrh range, Utah. Cotypes.—U.S.N.M. nos. 15456, 15459. Glossopleura parabola (Hall and Whitfield) Ogygia parabola Hall and Whitfield, U. S. Geol. Expl. goth Parallel, vol. 4, p. 245, pl. 2, fig. 35, 1877. Bathyuriscus productus Walcott (part), U. S. Geol. Surv. Bull. 30, p. 217, pl. 30, fig. 1e only, 1886. (See G. producta.) Dolichometopus productus Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 360, pl. 53, figs. 2d only (copy of original figure), 1916. This is a good species and should not have been included with G. producta. Occurrence same as preceding. Holotype—U.S.N.M. no. 15456d. Glossopleura packi, n. sp. Bathyuriscus productus Walcott (part), U. S. Geol. Surv. Bull. 30, p. 217, pl. 30, figs. 1c, Id only, 1886. (See G. producta.) Figures copied in Smithsonian Misc. Coll., vol. 64, no. 5, pl. 53, figs. 3-3, 1916. Bathyuriscus productus Pack, Journ. Geol., vol. 14, p. 207, pl. 2, figs. 3-3b, 1906. (Also 4b which he assigned to Alokistocare piochensis. ) The cranidium of this species differs from G. producta in having slightly smaller eyes and a wider glabella. On the other hand the pygidium of G. packi is close to G. parabola and if we had the cephalon of the latter, it also might be very similar. From G. parabola, the new species differs chiefly in a more circular outline, a slightly wider axis, and a relatively narrower doublure. Middle Cambrian, Chisholm ; (loc. 31) 3 miles northwest of Pioche, Nevada. Holotype and paratypes—U.S.N.M. no. 15455. Glossopleura belesis (Walcott) Bathyuriscus belesis Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 338, pl. 50, figs. I-17, 1916. Middle Cambrian, Gordon; (loc. 4v) 6 miles up Gordon Creek, Lewis and Clarke County, Montana. Cotypes—U.S.N.M. nos. 62667-76. Glossopleura bion (Walcott) Dolichometopus bion Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, pl. 52, figs. 2-2c, 1916. There is some hesitation to placing this form in Glossopleura, but the pygidium is altogether typical as are also the eyes and palpebral lobes. The difference lies in the slightly concave brim and expand- ing glabella. NO. 5 CAMBRIAN TRILOBITES—RESSER 33 Middle Cambrian, Spence; (loc. 55c) Liberty Canyon, west of Montpelier, Idaho. Cotypes.—U.S.N.M. nos. 62709-62712. Glossopleura utahensis, n. sp. Bathyuriscus productus Walcott (part), U. S. Geol. Surv. Bull. 30, p. 217, pl. 30, figs. 1, 1b, 1886. Bathyuriscus anax Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 335, pl. 48, fig. 1b, 1916. Besides the figured head and cheek numerous other cranidia as well as pygidia are on hand. Middle Cambrian, Ophir ; (loc. 30a) 1 mile below Argenta, Cotton- wood Creek, Wasatch Range, Utah. Holotype and paratypes——U.S.N.M. no. 62641. Glossopleura mckeei, n. sp. Dolichometopus productus Walcott (not Hall and Whitfield), Smithsonian Misc. Coll., vol. 64, no. 5, p. 360, pl. 53, figs. 4, 4a, 1916. Dolichometopus tontoensis Walcott (part), idem, pl. 51, figs. 1d’, 1d”, th (others Anoria tontoensis). Compared with G. producta, this species has a wider glabella, which expands forward somewhat faster and the pygidium lacks the pleural furrows and has much weaker axial furrows. In this respect it is more like G. parabola. The specific name is in recognition of the splendid work the present Park Naturalist Edwin D. McKee is doing at the Grand Canyon. Middle Cambrian, Bright Angel; (loc. 74e) Indian Garden Springs and (loc. 74) Nankoweap Basin, Grand Canyon, Arizona. Glossopleura buttsi, n. sp. Dolichometopus ? productus Butts (not Hall and Whitfield), Geol. Surv. Ala- bama, Spec. Rep. 14, pl. 5, figs. 8, 12, 17, 18, 1926. This species differs from G. producta in having shorter cranidium and pygidium. Middle Cambrian, Conasauga; Aldrich, Alabama. Cotypes—U.S.N.M. no. 90169. Glossopleura alabamensis, n. sp. Dolichometopus ? productus Butts, Geol. Surv. Alabama, Spec. Rep. 14, pl. 5, figs. 6, 7, 10, II, 13-16, 1926. Compared with G. buttsi, this species has a smaller eye and less fusion of the pleura in the tail. Middle Cambrian, Conasauga ; 3 miles west of Talladegea, Alabama. Cotypes—U.S.N.M. no. 90170. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Glossopleura lodensis (Clark) Bathyuriscus howelli lodensis Clark, Univ. California Publ., Bull. Dep. Geol Sci., vol. 13, p. 6, 1921. Dolichometopus ? lodensis Resser, Smithsonian Misc. Coll., vol. 81, no. 2, p. 10, pl. 3, fig. 9, 1928. Middle Cambrian; Marble Mountains, Mohave Desert, California. Holotype—vU.S.N.M. no. 78400. Glossopleura mohavensis, n. sp. Dolichometopus ? productus Resser, Smithsonian Misc. Coll., vol. 81, no, 2, p. 10, pl. 3, fig. 9, 1928. Occurrence same as preceding. Holotype-—On U.S.N.M. no. 78400. GLYPHASPIS Poulsen, 1927 Glyphaspis Poulsen, Meddels. Gr¢nland, vol. 70, p. 273, 1927. Poulsen described a species from Greenland, which may have to be referred to another genus. Comparisons.—As stated by Poulsen this genus differs from Asa- phiscus in having genal spines, long-pointed pleura, and a broad, fur- rowed pygidium, the pleural furrows of which are continued into the concave border. Genotype —Asaphiscus? capella Walcott. Range.—Middle Cambrian. DESCRIBED SPECIES REFERRED TO GLYPHASPIS Glyphaspis capella (Walcott) Genotype Asaphiscus ? capella Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 385, pl. 59, figs. 2-2c, 1916. Glyphaspis capella Poulsen, Meddels. Gr¢gnland, vol. 70, p. 273, 1927. Middle Cambrian; (loc. 54z) Half Moon Pass, Big Snowy Moun- tains, Montana. Cotypes—U.S.N.M. nos. 62761-4. Glyphaspis calenus (Walcott) Asaphiscus calenus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 384, pl. 60, figs. I-1c, 1916. Middle Cambrian, Meagher ; (loc. 5£) 11 miles south of Neihart, and Dry Creek, East Gallatin River, Montana. Cotypes—U.S.N.M. nos. 62765-62768. Glyphaspis camma (Walcott) Asaphiscus camma Walcott, Smithsonian Misc, Coll., vol. 64, no. 5, p. 384, pl. 60, figs. 2-2c, 1916. NO. 5 CAMBRIAN TRILOBITES—RESSER 35 Middle Cambrian, Meagher ; (loc. 4g) 5 miles northeast of Logan, Montana. Cotypes—U.S.N.M. nos. 62769-62772. Glyphaspis ? montanensis (Whitfield) Crepicephalus (Loganellus) montanensis Whitfield, Ludlow’s Rep. Recon. Yellowstone Nat. Park, War Dep., p. 141, pl. 1, figs. 1, 2, 1876. Specimen too poor to be sure of generic position. Middle Cambrian, Meagher ; near Fort Logan, 18 miles northwest of White Sulphur Springs, Montana. Holotype—U.S.N.M. no. 90668. IDAHOIA WALCOTT, 1924 Idahoia Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 58, 1924; idem, no. 3, Pp. 94, 1925. Genotype.—I. serapio Walcott. Range.—Middle Upper Cambrian. ADDITIONAL SPECIES REFERRED TO IDAHOIA Idahoia wisconsinensis (Hall) Conocephalites wisconsinensis Hall, 16th Ann. Rep. New York State Cab. Nat. Hist., p. 164, pl. 7, figs. 39, 41; pl. 8, figs. 22-24, 27, 28, 1863. Saratogia wisconsinensis Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 198, pl. 34, figs. 5-5c, 1916. Upper Cambrian, Franconia; Trempealeau and other localities in Wisconsin. Cotypes—A.M.N.H. Idahoia latifrons (Hall) Conocephalites latifrons Hall, 16th Ann. Rep. New York State Cab. Nat. Hist., p. 122, pl. 7, fig. 40, 1863. This form is a good species. Upper Cambrian, Franconia ; Trempealeau, Wisconsin. Holotype.—Presumably lost. Idahoia hamulus (Owen) Lonchocephalus hamulus Owen, Rep. Geol. Surv. Wisconsin, Iowa, Minne- sota, p. 576, pl. 1A, figs. 8, 12, 1852. Conocephalites hamulus Hall, 16th Ann. Rep. New York State Cab. Nat. Hist., p. 166, pl. 7, figs. 43, 44; pl. 8, figs. 25, 26, 1863. Saratogia hamulus Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 196, Ig16. Upper Cambrian, Franconia; Minneiska, Minnesota; Trempealeau and other localities in Wisconsin. Cotypes.—Original types destroyed ; plesiotypes, A.M.N.H. no. 315. 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Idahoia hera (Walcott) Saratogia hera Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 197, pl. 35, figs. 3-3b, 1916. Upper Cambrian, Franconia; Marine Mills, Minnesota. Cotypes—U.S.N.M. nos. 61715-6. INGLEFIELDIA Poulsen, 1927 Inglefieldia Poulsen, Meddels. Gr¢gnland, vol. 70, p. 261, 1927. This genus is closely allied to Kochiella, differing chiefly in having a wider rim and lacking the doublure impression. The rim in turn has the backward expansion opposite the center of the glabella. Pygidium unknown. Genotype.—TI. porosa Poulsen. Range.—Upper portion of Lower Cambrian. At present no Middle Cambrian species have been determined. Inglefieldia porosa Poulsen scribed as Chancia Inglefieldia planilimbata Poulsen Inglefieldia thia Poulsen (not Wal- Inglefieldia groenlandica Poulsen cott’s species ) Inglefieldia inconspicua Poulsen Inglefieldia affinis Poulsen = new Inglefieldia discreta Poulsen genus. Inglefieldia venulosa (Poulsen), de- KOCHASPIS, n. gen. Numerous species referred to several genera, particularly to Crepi- cephalus, when the pygidia were known, appear to be closely allied to K ochiella. Diagnosis —Cranidium wide. Glabella wide, tapering, clearly de- fined, with two or more pairs of furrows. Brim wide, divided into two subequal parts by anterior furrow which is shallow in the center ; rim clearly defined. Fixed cheeks two-thirds width of glabella; eye lines heavy; eyes rather small, situated at about the mid point of cranidium. Free cheeks extended into long genal spines. Pygidium with large prominent axis. Pleural furrows strong ; pleural lobes drawn out into spines. Surface granulose, lined or both. Genotype.—Crepicephalus liliana Walcott. Range.—Late Lower and Middle Cambrian. DESCRIBED SPECIES REFERRED TO KOCHASPIS Kochaspis liliana (Walcott) Crepicephalus liliana Walcott (part), U. S. Geol. Surv. 30, p. 207, pl. 28, figs. 3, 3a, 1886. (Not 3b, 3c—K. highlandensis) ; toth Ann. Rep. U. S. Geol. Surv., p. 653, pl. 96, figs. 7, 7a, 1890; Smithsonian Misc. Coll., vol. 64, no. 3, p. 209, pl. 29, figs. 5, 5a, 1916. NO. 5 CAMBRIAN TRILOBITES—RESSER By, Careful separation of the specimens identified as this and other species according to locality necessitates a realignment. Lower Cambrian, Pioche; (loc. 31a) Panaca Road, southeast of Pioche, Nevada. Cotypes—U.S.N.M. no. 15428. Kochaspis augusta (Walcott) Crepicephalus augusta Walcott (part), U. S. Geol. Surv. Bull. 30, p. 208, pl. 28, fig. 2a, 1886. (Not 2=K. nevadensis; 2b = another genus.) Locality same as preceding. Holotype-—U.S.N.M. no. 15430. Kochaspis nevadensis, n. sp. Crepicephalus augusta Walcott (part), U. S. Geol. Surv. Bull. 30, p. 207, pl. 28, fig. 2, 1886. (See preceding. ) Published figures exaggerate the narrowness which characterizes this species. Lower Cambrian, Pioche formation; (loc. 30) 8 miles north of Bennetts Spring, Highland Range, Nevada. Holotype —U.S.N.M. no. 61643. Kochaspis highlandensis, n. sp. Crepicephalus liliana Walcott (part), U. S. Geol. Surv. Bull. 30, p. 208, pl. 28, figs. 3b, 3c, 1886. (See preceding.) Rim on published figure drawn too concave. Pygidium like K. augusta but with narrower axis. Occurrence same as preceding. Cotypes—U.S.N.M. nos. 61640-1. Kochaspis cecinna (Walcott) Crepicephalus cecinna Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 99, Ok Tits WER, My i, OZ Lower Cambrian, Mount Whyte; (loc. 63a) Ptarmigan Peak, 54 miles northeast of Lake Louise, Alberta. Cotypes—U.S.N.M. nos. 64365-6. Kochaspis celer (Walcott) Crepicephalus celer Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 101, pl. 11, fig. 2, 1917. Ptychoparia clusia Walcott, idem, pl. 11, fig. 3. Middle Cambrian, Ptarmigan; (loc. 58k) Mount Stephen, 3 miles east of Field, British Columbia. Holotype —U.S.N.M. no. 64367 ; paratype, no. 64368. Kochaspis chares (Walcott) Crepicephalus chares Walcott, Smithsonian Misc. Coll., vol. 67, no. 2, p. 35, pl. 6, figs. 5-5c, 1917. 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Cranidium may belong to another species. Middle Cambrian, Ptarmigan ; (loc. 63d) east base Ptarmigan Peak, 3 miles northeast of Lake Louise, Alberta. Cotypes.—U.S.N.M. nos. 63744-6. Kochaspis ? gogensis (Walcott) Ptychoparia gogensis Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 88, pl. 12, figs. 4, 4a, 1917. Lower Cambrian, Mount Whyte; (loc. 62w) above Gog Lake, Wonder Pass, British Columbia. Holotype-—U.S.N.M. no. 64386. Kochaspis carina (Walcott) Ptychoparia carina Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 80, pl. 13, fig. 6, 6a, 1917. Middle Cambrian, Ptarmigan ; (loc. 35m) 3 miles southwest of head of Lake Louise, Alberta. Holotype —U.S.N.M. no. 64400. ,Kochaspis upis (Walcott) Crepicephalus upis Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, p. 218, pl. 33, figs. 4-4d, 1916. Middle Cambrian, Gordon Mountain; (loc. 150b) Chinese Wall, South Fork, White River-Indian Creek Pass, Montana. Lectotype-—U.S.N.M. no. 61697; paratypes, nos. 61695, 61696, 61608. Kochaspis unzia (Walcott) Crepicephalus unzia Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 3, p. 217, pl. 34, fig. 7a, 1916. (Not 7= undescribed genus.) Occurrence same as preceding. Holotype —U.S.N.M. no. 61706. KOCHIELLA Poulsen, 1927 Kochiella Poulsen, Meddels. Grgnland, vol. 70, p. 259, 1927. We are here dealing with a group of trilobites which developed greatly in late Lower Cambrian time, continuing into the Middle Cambrian. Aside from the two genera made by Poulsen, this group has long needed complete revision. Even now the final word cannot be said. Diagnosis —Trilobites with wide cranidia. Glabella clearly defined, with three pairs of furrows usually developed. Brim wide flat or concave, with a narrow rim faintly outlined ; conspicuous feature of brim is impression of doublure on upper surface, which simulates a NO. 5 CAMBRIAN TRILOBITES—RESSER 39 wide rim bent back toward the glabella at the middle. Fixed cheeks wide, eye lines heavy ; eyes moderate in size and situated rather far back. Free cheeks greatly expanded at genal angles, flat or concave in the outer portion and with long, wide genal spines. Pygidium with wide axis. Pleural furrows well defined and pleural lobes extended into spines. Surface usually with scattered granules often of more than one size. Owing to the width of the fixed cheeks, some species were referred to Olenopsis, while those for which the pygidium was known were placed in Crepicephalus. Genotype.—K. tuberculata Poulsen. Range.—Later Lower Cambrian, extending into Middle Cambrian. DESCRIBED SPECIES REFERRED TO KOCHIELLA Besides the species discussed below, there are K. propinqua Poulsen, K. arcana Poulsen, and K. gracilis Poulsen. Kochiella tuberculata Poulsen Kochiella tuberculata Poulsen, Meddels. Grgnland, vol. 70, p. 259, pl. 15, figs. 7-13, 16, 1927. Crepicephalus cf. cecinna Poulsen, Meddels. Grgnland, vol. 70, p. 267, pl. 16, HOS TZ) 1027. There can be but little doubt that this pygidium represents a species of Kochiella. Lower Cambrian, Cape Kent; Cape Kent, North Greenland. Cotypes.—Min. Mus. Copenhagen. Kochiella crito (Walcott) Olenopsis crito Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 75, pl. 11, fig. 6-6b, 1917. Lower Cambrian, Mount Whyte; (loc. 60e) Ptarmigan Lake Pass, Alberta. Lectotype —U.S.N.M. no. 64371; paratypes, nos. 64372-3. KOCHINA, n. gen. Certain Middle Cambrian species resemble Kochiella, but all differ in the same manner and therefore are regarded as a distinct genus. Comparisons.—K ochina differs from Kochiella in three respects: First, the brim is narrower so that the preglabellar area is practi- cally eliminated ; second, the converging course of the anterior facial suture greatly reduces the area of brim at the anterior angles; and third, the eyes have a more anterior position. 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Genotype.—Olenopsis americanus Walcott. Range.—Thus far confined to the Albertella zone of the Middle Cambrian. DESCRIBED SPECIES REFERRED TO KOCHINA Kochina americana (Walcott) Olenopsis americanus Walcott, Smithsonian Misc. Coll., vol. 57, no. 8, p. 243, pl. 36, figs. 8-11, 1912. Middle Cambrian, Gordon; (loc. 4v) Gordon Creek, Montana. Cotypes.—U.S.N.M. nos. 58368-71. Kochina bosworthensis, n. sp. Olenopsis cf. americanus Walcott, Smithsonian Misc. Coll., vol. 67, no, 2, p. 37, pl. 6, fig. 8-8b, 1917. Middle Cambrian, Ptarmigan; Mount Bosworth (loc. 35c) and Popes Peak, above Ross Lake (loc. 63j), British Columbia. Lectotype —U.S.N.M. no. 63749; paratype, nos. 63450-1. MACELLOURA, n. gen. Diagnosis —Cranidium quadrate, facial sutures diverge anterior to eye, making sharp, salient, down-turned anterior angles; postero- lateral limbs short. Glabella quadrate, tapering but slightly ; extends to anterior margin; without glabellar furrows; wide. Fixed cheeks one-fourth width of glabella; eyes rather small. Pygidium spade-shaped with anterior pleural furrows making up- turned sides; dorsal furrow marked only by change in slope; axial and pleural lacking ; axis tapering rapidly, half as long as pygidium. Free cheeks and thorax not known. Genotype.—lllaenurus ? dia Walcott. Name.—Maked\a = spade ; ovpa= tail. Range.—About middle Upper Cambrian, or somewhat younger. Macelloura dia (Walcott) Illaenurus ? dia Walcott, Proc. U. S. Nat. Mus., vol. 13, p. 277, pl. 20, fig. 6, 1890. Nileus ? dia Walcott, Smithsonian Misc. Coll., vol. 57, no. 13, p. 359 (gen. ref.), I914. Upper Cambrian, Wilberns; (loc. 70a) Morgans Creek, Burnet County, Texas. Cotypes.—U.S.N.M. no. 23865. METEORASFPIS, n. gen. Diagnosis ——Cranidium but little larger than glabella. Glabella large ; dorsal furrow well defined ; without glabellar furrows. Occipital NO. 5 CAMBRIAN TRILOBITES—-RESSER 4! furrow and rim well developed. Brim about one-third as wide as glabella is long; heavy, thickened, up-turned rim separated by strong anterior furrow; preglabellar area almost obsolete in front of glabella. Fixed cheeks narrow, consisting entirely of the strongly arched palpe- bral lobes and apparently narrow straps. Eyes rather large. Genotype-—Ptychoparia ? metra Walcott. Range.—Middle Upper Cambrian. Name.—Mereopos = high ; aoms = shield. Meteoraspis metra (Walcott) Ptychoparia ? metra Walcott, Proc. U. S. Nat. Mus., vol. 13, p. 273, pl. 21, fig. 7, 1890. Upper Cambrian, Wilberns; (loc. 67) Potatotop, 7 miles northwest of Burnet and (loc. 68) Packsaddle Mountain, 11 miles southeast of Llano, Texas. Holotype —U.S.N.M. no. 23858. MODOCIA Walcott, 1924 Modocia Walcott, Smithsonian Misc. Coll., vol. 75, no. 2, p. 59, 1924; idem, no. 3, p. 106, 1925. Genotype—Arionellus (Crepicephalus) oweni Meek and Hayden. Range.—Upper Cambrian. DESCRIBED SPECIES REFERRED TO MODOCIA Modocia oweni (Meek and Hayden) Arionellus (Crepicephalus) oweni Meek and Hayden, Proc. Acad. Nat. Sci. Philadelphia, 1861, p. 436. Modocia oweni Walcott (part), Smithsonian Misc. Coll., vol. 75, no. 2, p. 59, pl 12) figs. 1, 2, 1924- idem, no. 3, p: 106, pl. 16, figs, I, 2, (not 3= WM. centralis), 1925. (Not Ptychoparia oweni Walcott = Ehmania owent. ) This trilobite was referred by various authors to Agraulos, Ptycho- paria, and Crepicephalus. Also several other species were included under this name. Study of the types shows that the species occurs only at the locality given below. Upper Cambrian, Deadwood; head of Powder River, Big Horn Mountains, Wyoming. Holotype —U.S.N.M. no. 1180. Modocia centralis (Whitfield) Crepicephalus (Loganellus) centralis Whitfield, Prelim. Rep. Pal. Black Hills, U. S. Geol. Surv., p. 10, 1877; Rep. Geol. Res. Black Hills, U. S. Geogr. and Geol. Surv., p. 341, pl. 2, figs. 21-24, 1880. Modocia owem Walcott (part), Smithsonian Misc. Coll., vol. 75, no. 2, p. 59, pl. 12, fig. 7, 1924; idem, no. 3, p. 106, pl. 16, fig. 3, 1925. 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Upper Cambrian, Deadwood; Castle Creek, Black Hills, South Dakota. Cotypes—U.S.N.M. no. 24581. Modocia berkeyi, n. sp. Agraulos convexus Berkey, Amer. Geologist, vol. 21, p. 288, pl. 20, figs. 9-11, pl. 21, figs. 3, 7, 1808. Agraulos convexus var. A. Berkey, idem, pl. 20, figs. 1, 2, pl. 21, fig. 5. Ptychoparia calymenoides Berkey, idem, pl. 20, figs. 3, 4, pl. 21, fig. 4. It appears that variety A and the other specimens assigned to the species are alike except for size. However, critical work will have to be done to settle this point. Variety B of Berkey belongs to another genus. Upper Cambrian, Ironton; Taylors Falls, Minnesota. Holotype-——Columbia Univ. no. 22283; paratypes, nos. 22286, 22307. Modocia glomerata (Walcott) Acrocephalites ? glomeratus Walcott, Smithsonian Misc. Coll., vol. 64, no. 3, Po l70, pl. 20, figs. 7, 7a; 1916: This specimen is typical of the genus Modpcia, departing only in the development of the median boss. Upper Cambrian, Deadwood ; (loc. 340c) near Rawlings, Wyoming. Holotype—U.S.N.M, no. 615901. Moosia Walcott is a synonym of Elvinia Walcott (Smithsonian. Misc. Coll., vol. 75, no. 3, p. 88, 1925). PLAGIURA, n. gen. Diagnosis —Cranidium large, smooth, rather flat; facial suture converges slightly anterior to eyes ; behind the eyes it diverges rapidly, forming wide, blunt postero-lateral limbs. Brim about one-third as wide as the glabella is long; no rim separated. Fixed cheeks about half width of glabella; eyes small, situated opposite anterior third of the glabella. Free cheeks simple curved bands, with rounded genal ‘angles. Pygidium triangular, broad, with anterior angles somewhat drawn out; axis wide, well separated by change in slope and extends nearly to rear margin ; axial and pleural furrows present. Genotype.—Ptychoparia? cercops Walcott. Range.—Upper Lower Cambrian, Canadian Rockies. Name.—rAaywos = slanting ; ovpa=tail. NO. 5 CAMBRIAN TRILOBITES—RESSER 43 Plagiura cercops (Walcott) Ptychoparia ? cercops Walcott, Smithsonian Misc. Coll., vol. 67, no. 3, p. 81, pl. 12, figs. 1-1d, 1917. Lower Cambrian, Mount Whyte; (loc. 63c) 14 miles northeast of Lake Louise, Alberta. Cotypes—U.S.N.M. nos. 64377-64381. POLIELLA Walcott, 1916 Poliella Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 340, 1916. This genus was established as a subgenus of Bathyuriscus, with the statement that the pygidium was different. Even a cursory revision of the trilobites referred to Bathyuriscus and related genera shows the necessity of making this a full genus. Evidently, Walcott first in- tended to use the form from Pole Creek, Bathyuriscus powersi, as the genotype, but since he designated the Spence shale form, it be- comes the genotype, and the genus is not represented at the Pole Creek locality. Diagnosis —Cephalon much larger than pygidium. Glabella quad- rate, expanded anteriorly; two pairs of glabellar furrows usually clearly defined, others very faint or absent. Eyes long, moderately bowed. Brim when present narrow and simple. Thorax with 7 to 9 segments. Pygidium with wide, well-defined axis; pleura usually fused ; two to four pleural furrows extend to the margin; rim absent. Comparisons——Compared with Bathyuriscus, Poliella is distin- guished by its fewer thoracic segments, larger eyes, and much smaller tail. Genotype-—Bathyuriscus (Poliella) anteros Walcott. Range.—Middle Cambrian. Species formerly referred to Poliella: P. powersi= Bathyuriscus P. sylla= Clavaspidella P. probus = Clavaspidella DESCRIBED SPECIES REFERRED TO POLIELLA Besides the species discussed below, the following belong to the genus. Poliella anteros Walcott Poliella chilo Walcott Polhiella balus Walcott 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Poliella occidentalis (Matthew) Dolichometopus occidentalis Matthew, Trans, Roy. Soc. Canada, 2d ser., vol. 5, sec. 4, p. 49, pl. 2, fig. 2, 1890. Bathyuriscus pupa Matthew, idem, p. 51, pl. 2, fig. 5. Bathyuriscus occidentalis Walcott, Smithsonian Misc. Coll., vol. 53, no. 2, p. 41, 1908. Walcott, Canadian Alpine Journ., vol. 1, pt. 2, pl. 3, fig. 2, 1908. Bathyuriscus (Poliella) occidentalis Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 351, pl. 46, fig. 3, 1916. Owing to the poor preservation of the holotype, this species would be inadequately represented except that two excellent carapaces have just been discovered in our collections. It seems that the pygidium fits Poliella. The published figures are restored more than warranted. Middle Cambrian, Stephen ; (loc. 14s) Mount Stephen, near Field, British Columbia. Holotype—Royal Ontario Museum (casts U.S.N.M. no. 62621). . Poliella prima (Walcott) Bathyuriscus (Poliella) primus Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 352, pl. 46, figs. 6, 6a, 6b, 1916. (Not 6c, d=P. castlensis.) This is described as a Lower Cambrian species but belongs to the Ptarmigan formation. Middle Cambrian, Ptarmigan; (locs. 35m, 35e) 3 miles southwest of the head of Lake Louise, Alberta. Lectotype-—U.S.N.M. no. 62624; paratype no. 62623. Poliella castlensis, n. sp. Bathyuriscus (Poliella) primus Walcott (part), Smithsonian Misc. Coll., vol. 64, no. 5, p. 352, pl. 46, figs. 6c, d, 1916. It is necessary to separate this head from P. prima because of its wider brim and eyes which have a more divergent position. Middle Cambrian, Ptarmigan ; (loc. 58t) Castle Mountain, Alberta. Holotype —U.S.N.M. no. 62626. Poliella caranus (Walcott) Bathyuriscus (Poliella) caranus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 350, pl. 46, fig. 5, 1916. This form barely remains within the genus because of its greatly reduced brim. Middle Cambrian, Spence; (loc. 55c) Liberty Canyon, west of Montpelier, Idaho. Holotype—U.S.N.M. no. 62628. —_ - NO. 5 CAMBRIAN TRILOBITES—RESSER 45 STROTOCEPHALUS, n. gen. Diagnosis —Cranidium alone available. Facial suture diverges an- teriorly to rather sharp anterior angles; glabella tapering with faint furrows ; occipital furrow shallow; occipital ring slightly thickened. Brim wide, concave, and without a rim, striated vertically ; eye lines developed; palpebral lobes of moderate size, situated far back. Comparisons.—Differs from Alokistocare in two points, viz, sharper anterior angles and more posterior position of the eyes. Genotype —S. gordonensis, n. sp. Name.—Xzporos = spread, xepados = head. Range.—At present confined to Middle Cambrian. Strotocephalus gordonensis, n. sp. Ptychoparia pylas Walcott (part), Smithsonian Misc. Coll., vol. 67, no. 2, pl. 6, fig. 4c, 1917. (Other figs. Elrathia.) Middle Cambrian, Gordon; (locs. 4q, 4v) Gordon and Youngs Creeks, Ovanda quadrangle, Montana. Holotype—U.S.N.M. no. 63743. WEEKSINA, n. gen. Two species from the Upper Cambrian of the House Range referred to the Middle Cambrian genus Asaphiscus are distinct in many respects. Diagnosis —Entire trilobite ovate; convexity not determinable. Facial suture diverges anterior to the eyes and is intramarginal two- thirds of the distance to the center so that the anterior angles are strongly rounded. Glabella wide, rounded in front; glabellar fur- rows short, but fairly deep. Brim of moderate width; rim thickened and, owing to course of the suture, sharply narrowed toward anterior angles. Fixed cheeks narrow; eyes fairly large; palpebral lobes strongly bowed. Free cheeks with well-defined rim and blunt genal angles. Thorax with 10 or 12 segments; tips of pleura blunt ; long spine on axis of next to last or third from last pleuron. Pygidium half the size of the cephalon; axis well defined, rather wide, and extending nearly to rear margin; axial and pleural furrows distinct. Surface granulose. Genotype.—Asaphiscus ? unispinus Walcott. Range.—Presumably early Upper Cambrian. 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS — VOL. 93 DESCRIBED SPECIES REFERRED TO WEEKSINA Weeksina unispina (Walcott) ‘ Asaphiscus ? unispinus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 389, pl. 61, fig. 1, 1916. Upper Cambrian, Weeks ; (loc. 30n) Weeks Canyon, House Range, Utah. Holotype —U.S.N.M. no. 62775. Weeksina granulata (Walcott) Asaphiscus ? granulatus Walcott, Smithsonian Misc. Coll., vol. 64, no. 5, p. 385, pl. 61, figs. 2, 2a, 1916. Number of thoracic segments uncertain. Occurrence same as preceding. Cotypes—U.S.N.M. nos. 62776-7. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 6 Pon OS LOSES (WitH Five PLATEs) BY ALES HRDLICKA Curator, Division of Physical Anthropology, U. S. National Museum (PUBLICATION 3296) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MAY 14, 1935 The Lord Baltimore Press BALTIMORE, MD., U. 8. Ay CONTENTS PAGE fra veC@ra ho Tek aya? fh as is teen prem ra ere Nie, RACE ECU ea TO ee he 1 I IL SHRED TREY ee are Rac Ri Creat OCS DI SEO TE HO ICE VO MEER EN ee I STITT TID CUIAV PA oA SeaR RNase hol yarate ea av aves ol Tek such hbase nh ICC NATE bie eeleveie 5 12 ald a Cee Go) 01 (=) ee ace ee a at He ee RS te 13 Minictacasas a (Chas) sete epee eres cer Sd more ee c Nulstele ics Mon treet atid crooks 14 Grier ALACES me (LADLE) Osc. noe cicee & octane aa oe chad ahs A ee ke ee 15 CO irrical EMA LeL tale Se cette otayehs a Io Wawreee rascal aia Ms Re ae Sea ee 16 INGqUODSGtava ELOUS Me eee eae ERIN Se CA Sea oT aR eae 19 INewarta terials eae ewes hey: Selotar eae Leyes. Cede ans cre SRO RENE ate 20 [Dysitenii bevel) GER ares Br Sener eave tear ens a Eitan His mene ea gE NE Bm GEr ree PAAERL aes tae 23 INGO Poh CIC TCR ETRE ECE TS CIE OOS OI Ona MCI Chon ueee CI SIOe 23 Sree td atte ed Se SIN ene En ert te i oer Rta ie BU I eae Ca 27 Breqnency, of exostoses) in the two eansees ass ss sce aaate elon eiieiciele 28 SHAS 5) S55 Oe OOO Orig SEITE OES Cie Seon ce ena eR eae As a me, oat 29 INGittTa De Seas hele tavaee da sereTe are te Sea I Oe oe ec aes dk Ht ag ee, 31 LE YOSSI VO beer tear aR Rcetak sca RRR ERE eee nee MP eS NENA CRI, Bd 33 SEAS on BE ek eae as Br A a fe Fn OR Se SC RR er 8g ae M&O By) SEEN DYES wis rs Seno care CEC CE MCnar tC Re Ts CIPO IT er CRE ea a Un en IE 5 Se A 39 SATHMLEL TVR eeercrere tere Sie coe tir ceca RTT ea ere a are ee ah aye 40 Ate aRO LE SRO Withicnes a asec hal ssts rier era ee eo creas re are Oe 4I SHULoxa bi soTesWosal Sp hea ann Ne Be co Te en Po Ore Mm oa tere 1) i 42 SEMUICHITepiter rreNe ects eas reer chine Ina Pr cee nar aay eo HAE Me et Sate 46 |B SE te a Uae ee ney Sy sk EN UN Othe oa pn a ORR RO Ao a ata mae Rd 47 SENSOR Ee ere ea rare Tecate 67 arabia a relia EDS fouatd Seeye sigs 0, vide alas ay aha, eaets 49 AMER CATISES San LINC OWills Prattys are mhaese a ciet tas cine nates, ce ctiaceie eee te ever: 49 STORE RTC IN IS oh INTE TED VC? ocrt tsa.) la taltvin td aisinrsy Sere swe srerecaiasetets cei sina rene 50 (CHO eYERSTINGET AT apie crenetice ty Scien cic RT een CoE rare ee ee a ao a 50 Developiieritall 4, site avs/sasvel chavveestersieyeneitetereiet jars = ole rer er evens ee Stsye.elevane dh avetevecs SI iTBOLMtENMeN Cys, scltee aeie eos Se Mero aortic Se OUR clalinsiesiea am cele 53 let Citta tercretars cot cress hotel Pax eestor aay ere ERTS To eval uA eee GIN recA relied 54 Socialstattisi © ceupatton--/eaicncinecsiey eestor eres Sate, Ne nicer ovale: 57 @littrvet Cape rer ere arcs real oie eee es eg te ee cee SO ROL eI 58 SMM pe ett cae CREME tra ver Acct Nace javatrsley 5:0) exeiakats fier deatests RAWiels Sf Seerea days lave a hohe 58 Constiutionaladerancementse, q.coeeee eee cmeon een ceen cea 59 Goutsparthritd cadiathests)er sts sissies cree Sethe eye en a oes 59 SS Uypia MUL Tem agora teen eye cin Seder ohtuaerae sere een Ais cea Aa oS soda NS, aT 61 ocal igriwations and cintlammatiOn 5.50 saice ayn. oiciad etcia's'e 6 se, sc 0/o.e wid ae! ate breve 62 SEU SPEIER a SRSA nea 2 vee get ek OS CE Ce ee en 65 JASISOSSINEIS: A TITGEIIYG Fete CR mesma WN ea ne a Pr en ee 66 aE MN reer esse ies AD ap Yoh maybe Ass eee choke eae che 0: 6 nants bevel 66 Wiechat cal ncalisessarid framiatisil acc a n.-o,aoe A Ta os era ceo ke G owaied oe 66 EI eAM Pe LOniA LION cratyonen «tie othe Namie ok Suis ck ooo oA cole dena lee bos 68 iv CONTENTS VOL. 93 PAGE Miscelllatieotis’ \CatS@Si asc orescorec ters acs.o a innover ovate e havatetee enerojee tenn teuerrens Celtel 73 Enbineedimeri ocr otic fee icrectes nh dis2 oie joieathe eta a ata neers eae vet eater 73 Constitutional diseases (other than syphilis and arthritis)......... 73 Trim @ Verb: eh reccsoce gc ch cue ccova bos Palo ates 0 on orwe lets hee aovetonner cacteseieietenavene te eral 74 Connection with exostoses of inner ear, or of other parts of the CCA fol) ee eS et oe Giant OSG OO GOODS ULI 74 OtOSClERGSIS= bac cindc.cs 5 isc nie wittleteciane etre eee oer eer traits 74 NY, Ft 10) (a ee ER RIE IR cin eutas Cm GRO CLAS Serco CL OOS 74 INAa Stat LOMI cote s:cis, eiigrovs oy o.sichesd lesa e Outlet toser ene eee SIG nee eLaTe oe rere 74 WeStistall So cincdscies,s as a. covam ortattierer tactayesiay hab aciepeletn tae aan 75 INO path OlOpy= c= s.« jcreie «.s: serene wi oiccayo nya:elinleiatienetepal ne onataye cheep unten amare 75 Genekal DIScussion. '5.0e.-**yJassny | O061 Paine dnoiry ioyyny Iva (0) OI! “-- “mbeyoyey 0'9 + oS miewne aoteiel aren nia ‘aqRy ud 9681 Onl 9 OOr a Sc eae nid re Peete KORIOG] +6g1 Tod I 16 eA] Hee ome uueuys— F6gI gor oe ni fetes tt naag bv &T QI PEI | *(uooUY) nNiag |° °°" MOYoITA Cggi g If Z (AGM Non aae UMal(dooy- ||P 272" Aaypnys Iggl 9 . b c gol LO et hit eda) yyeg S° 91 gt C61 UNS |p axPI1G Oggl ie . 9 6 Lt Coot en Pe CR ier niag aoe z ¢ uua I'6 I Il 208 6 PBL TE(g{ []o oo 2 925 JIMO]] 6Lg1 . : “++ 55+syoouryD I d Paster) Siete Pat ets niag eho n baie JauUIN [, 6Zg1 bv €1 6 L9 EUS fee, eae die) gLgI be~2 g ODES ee eas: WB G>s ele oP oe uewA\\ tlegl tv é wi leJlettayie) selma e nidg PRPS OP SUNG ¢Z-LogI I 5 @ 0 6 6 inte pu] xO 5) ome ues “IDYOA\\ tog C 9 bi ieniene Helvetie:p 5 nidg am eke UUPLUSI[IS togi quaciag Resaes ee. dnoiy 1oyyny 1e9aX sqnys | ~ UDIPUT UDILLIU PY AY4 UL Sasojsoxy 4D uo DIDG fo aunsay *QINjXIUIPe URISOURTaW euOSs A[qeqoid yW1M 4Nq ‘uUeISeUATOd vo £ 9—z—tz—,,purys] Josey NO. | | _ UOUIWIOD, , é | ¢ é < HEA S > a 1'6 | 17 | tr 3 spurys] weyzey) ma £°9 | I | ZI a “‘YoIy Joqwuius T Z | rs oO é z é I é ty) to) 16+ e ! é Ny spurs] AjeAo7] ODRISy,, fo) mA a) Loz t6'o I Lor or | z | oz | € é | I | : SUPISOULLITL spurys] sesonbiey, jedan % os i A a i ee Sasoy sasoy saso} soso} ot -soxa -sOxe -soxa oi -soxa qusd10g ava s]INyS | qued10g iva s][NyS | JusdIeg Bich) s][INyS | Jued1eg 1a S]INAS quoad1eg iea SUDAS yam yam ya ae yan sIINAS sIIMyAS SIINAS SIINAS (O139N) BOTW eIPesqjsny BISOUL]AIL BISoUA|Og PISy DILIU Y PUD adOANG] UDY] 40410 SUOIBIY U1 SasOjSOXT AD \o 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 The maximum incidence of ear exostoses is not in Peru, but ap- parently among the tribes with fronto-occipital head deformation of the Columbia watershed, and in the Tennessee-Ohio mound and stone- grave region; also apparently in the old population that has left its skeletal remains in the caves of Coahuila, northeastern Mexico. The data will receive further attention in the discussion of causa- tion. For the present it is sufficient to note the fact that in America itself very marked differences exist, as to tribe and location, in the frequency of the growths under consideration. There is but a single observation on a series of Eskimo skulls and that is negative—Russell in 54 specimens found no exostosis. From regions other than Europe and America the data on ear ex- ostoses are as yet very inadequate. There is little or nothing in this respect on the peoples of the densely populated areas of China, India, and Malaysia, of the rest of Asia or of Negro Africa, but there are indications that the abnormalities in question are relatively frequent among the Polynesians. The data are given in the table on page 15. CLINICAL MATERIAL In addition to the data tabulated in the preceding pages, there are scattered through medical literature of the last and present centuries reports of perhaps as many as 300 separate clinical cases of ear ex- ostoses in white Europeans and Americans. A large majority of these are reported more or less defectively as to nationality, sex, age, and other details of importance, attention having been centered on the pathology and cure of the condition. Anthropologically, they add but little to the knowledge of the growths beyond showing further that they occur with no great rarity in England, France, Belgium, Germany, Italy, and other European countries, as well as in the United States. Numerous as these reports are, they would not suffice to give a clear and full picture of the abnormalities. But here and there interesting points appear, and collectively there is much of value on the histology and especially on the etiology of these ear tumors. The earliest comprehensive account of this nature is that of Joseph Toynbee (1850). He reports 12 cases of such “tumors”, 10 (ap- parently) in males and 2 in females, mostly elderly people of British extraction. The exostoses consist of very hard and dense bone, are of slow growth, and develop frequently “ unattended with any symp- toms.” He strongly suspects their connection with gouty diathesis and from a further standpoint of causation divides them into two classes—one “ in which the disease appears associated with congestion No. 6 EAR EXOSTOSES—HRDLICKA Ly of the mucous membrane of the ear, as a result of rather free living ”’ (and drinking) ; and the second associated with “ disease in the cavi- ties containing the expansion of the auditory nerve.” The first “ thesis” devoted entirely to the subject of ear exostoses is by Delstanche (1878) ; and in the course of time this is followed by other such treatises by Schlomka (1891), Braunberger (1896), Sabroux (1901), and Bachauer (1909) ; to which may be added the comprehensive dissertations on the subject by Alexander (1930) and Moller-Holst (1932). It would seem that a relatively limited condition that has received such prolonged and extensive attention should be well understood by this time; but, as will be shown later, this is still far from being true. The records of over 200 cases of ear exostoses reported by aural surgeons are given in the next table. The stated nationality is usually that of the author, but the subjects, especially in such complexes as Austria or the United States of America doubtless included various other racial elements. The data as to the sex, side, etc., do not always extend to the whole or an equal number of cases reported, which reduces their value. They need not be considered at this juncture; as far as they go they will be dealt with in subsequent chapters. The table on page 18 has no pretense to completeness. There are records of additional cases, but some of these are little more than mere mentions, while with others the original report was unobtainable. Burkner (1884, pp. 92-93) summarized the statistical data on ear diseases published by 16 authors to 1884, but combined, regrettably, stenoses, exostoses, and atresiae into one group, and there is no way of separating the three. The figures had nevertheless some interest. In all there were 94 cases of the three above-named conditions, re- ported by 6 authors; and their combined frequency was, for Gruber 0.2; Lucae 0.2; Hedinger 0.3; Newark 0.3; Burkhardt-Merian 0.5; and Schwartze 0.2 percent of all cases with ear diseases. What stands out from all the preceding data is the wide distribution of the incidence of ear exostoses, geographically, racially, and also in time. It is no recent or local affliction, but an old and widely gen- eralized disorder. A few racial groups appear immune, but from none of these is there sufficient material to decide the question. In the remainder the frequency varies greatly, and this variance seems to follow racial or geographical affinities, but there appear no few exceptions to this, especially in the American Indian, where group conditions evidently prevail over the racial. 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Owing to some fortunate circumstances and with the invaluable aid of the Peruvian authorities, he was able to make large skeletal collections which, aside from a mass of other parts of the skeleton, comprised approximately 4,000 skulls (Hrdlicka, t911, 1914). These proceeded mainly from the coasts but in an important degree also from the moun- tains. A portion of this material was used in 1915 in the preparation of the anthropological exhibit for the Panama-California Exposition and remains in the San Diego Museum; the rest is preserved in the National Museum at Washington. Even in the field I was struck by the frequent presence in the outer half of the external auditory meatus of bony swellings and in some cases distinct tumors that more or less restricted or even almost closed the ear opening. In the report on the second expedition (Hrdlicka, 1914), I briefly called attention to the fact that “a rela- tively large proportion of the pre-Columbian people of the more cen- tral parts of the Peruvian coast suffered from a greater or lesser occlusion of the external auditory canals by bony tumors.” A small series of these crania with ear exostoses was exhibited and remained at San Diego, and another series was placed on exhibit in the Divi- sion of Physical Anthropology at the United States National Museum for demonstration to visiting surgeons. In 1921-22, at my instigation, the series of Peruvian skulls in the United States National Museum was examined for the bony tumors in question by Dr. Beatrice Bickel. During these examinations and especially when an attempt was made to prepare the results for publi- cation, it was found that the subject was more complex than at first anticipated and that some needed data were not obtained, which led to a second study of the whole series by Dr. Paul Van Natta, then Aid in the Division of Physical Anthropology of the Museum; and as the results of this second study differed somewhat from those of the first, a larger part of the collection was submitted to a third ex- amination by myself. The results differed slightly from those of both previous examinations, but the differences in the three were seen now to be mostly only those in the precise appraisal of the various grades of the abnormality, the essentials remaining fairly constant. Subsequently, finding the condition repeatedly also in other Indian and even in some non-American skulls, I extended the personal ex- amination to a number of series of crania in the National Museum collections. Then other work intervened, and the publication of the results had to be postponed. 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Meanwhile, as already noted in the first section of this memoir, the material left in San Diego received attention by a number of students from the west coast, two of whom, Burton and Moodie, reported on the ear exostoses of the skulls. Other publications on the growths appeared from time to time. From 1930 to 1932 three especially important studies in this line were published, namely those by Oetteking, Alexander, and Moller-Holst. Moreover, additions were received to our collections which promised to throw additional light on the peculiar disorder. These considerations led me to a de- termination to finish the survey of the subject, add as much as pos- sible to the previous data, and attempt, should the facts warrant, to advance the understanding of the affliction under consideration. The results follow. NEW MATERIAL The total number of skulls examined for ear exostoses in this new study is 7,814. All these, with the exception of about one-third of the Peruvians, were sexed and examined by me personally. The material is part of that of the Division of Physical Anthro- pology, United States National Museum, and the only portion of the specimens reported upon previously were those mentioned casually in my own publications. Only the larger series of our collections were made use of, and such as would give as far as possible a widespread racial as well as geographical and time distribution. The abnormal bone formations were found to range without any line of separation from distinct localized “ pearls” “ or tumefactions of bone in the external auditory canal to bony tumors that almost fill the distal part of the canal or even protrude outside of it. Only those cases were recorded where the growth presented distinct local- ized welting, excrescence, or tumor. In addition there were fairly numerous cases where a more or less evident trace of a tumefaction or a diffuse pathological thickening of the wall existed; these were not included in the records. The earlier and again the late stages of ear exostoses present diff- culties to the examiner. The initial tumescence, “ pearl” or welt, may be ill defined; and one or two small swellings, welts or “ pimples ”’ or ridges, may accompany a larger growth which tends to preempt the attention or obstruct clear vision. Hence no two observers or even repeated examinations by the same student will give absolutely ‘ * The term “pearl” is particularly fitting, for many of the growths in their earlier stages very closely resemble developing pearls on their mother shell. NO. 6 EAR EXOSTOSES—HRDLICKA 21 the same results as to the minor grades of the abnormalities. Consid- erable experience must be acquired by the student of these exostoses before his records can have real value. The total new material and the gross results of the examination are shown in the following table. Ear Exostoses in Different Racial Groups, New Observations NGBeE Number Percent Number Percent Adults ¢ F Bheulls of skulls | of skulls of ears of ears (except as specified otherwise) examined with ear with with with exostoses | exostoses | exostoses | exostoses Egypt: XG) eyleishitu 92 eeysesorreras 379 7 Tas, 10 Te? XX D.—III Cont. Kharga... 75 2 DoF 2 Tes American: | EHS) cer 6 (ee ae a I ,000 2 0.2 4 0.2 Indian @hildren®= see 335 I I hee Gi foT 21S] 0) Poa een re 500 12 24 19 1.9 Horta DakGtas i: ss. c55- 26 29 2 6.9 2 3-45 RilOniGda so eters cys oy acnaie si aarsie 395 35 8.9 53 6.7 Galifornmatn er (a sons cee 435 46 10.6 69 7.9 NE. States (St. Lawrence River to Maryland)....... 112 13 iE (0) 19 8.5 Greer red anicus een seerasie eid tcseose 3,651 522 14.3 855 TAT MUMIA Ae ec Ss wali 65 14 Bile fy 25 19.2 Louie, a5 e050 oco gp omaor 61 15 24.0 26 BN, IXTCATISASSernih ees is seit ore les 73 47 yee 75 2 Tiny, S. Dakota (All, Mowbridge, Arikan eeiniSGs) weiss eh qe 109 30 27.5 48 22.0 S. Dakota (Mowbridge alone) 76 23 30.3 37 24.3 INentuckymerrn ccs eee go 29 Bene 52 28.9 Polynesian: INewaZealandi.s5 cates ee 19 4 Bit sit 7 18.4 IS EN TANDI AA Setcige Mele Goole aide 148 39 26.4 60 20.3 Asiatic: Whinese ae Cee eessesrinds exe Wid Malaysian: Be AG lees aiasy sis. Shaverdio diel silers 10 Melanesian tote cctat ote. 39 African: Negro (African and American) I12 @ Including a very small proportion of subadults. + From age when fully developed tympanic bone is in place to eruption of a permanent second molar. A fair proportion of deformed, all three varieties of deformation. © Mainly of Channel Islands. 4 Mounds. Examination of our Egyptian material, all of which was collected personally and is thoroughly identified, shows that although ear ex- ostoses were rare, nevertheless they occured in the Egyptians—at root a branch of the White stem—as far back as 2000 B. C. i) i) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 There is no case of a tympanic exostosis in the 112 African and American Negro crania in our collection. It will be recalled that Ost- mann, in 267 Negro skulls, also found no case of these exostoses. It would seem, therefore, that the true Negro was free of these formations. No case of these tumors is found in our Melanesian and Malaysian skulls, but the series of specimens here are too small. Flower, it was seen, found one case of these exostoses in 1 of 7 Australian and in 1 of 20 Melanesian skulls. It appears, therefore, that the condition is not wholly absent either in Melanesia or Australia, though it prob- ably is rare in these territories. Especially interesting is the apparently complete absence of ear exostoses in the Chinese. No case in these people, so far as I could discover, is on record, and the valuable series of 77 adult male Can- tonese skulls in our collection shows not a trace of bony ear tumors. Yet the Chinese belong fundamentally to the same yellow-brown human stem as do the American Indians, who show so many of these abnormalities, while on the other hand they are far apart from the African Negro, who similarly appears to be free from ear exostoses. In the remaining groups of our table conditions differ greatly. At one end of the series stand the Eskimo, first cousins of the Indians, who in 1,000 skulls give but 2 with a moderate form of distinct local- ized tympanic tumefaction ; at the other end are the Kentucky, South Dakota, Arkansas, Louisiana, and Virginia Indians, with the Hawaii and New Zealand Polynesians, among whom over one-fifth to one-third of the skulls show the neoplasms under consideration. The Arkansas and Louisiana Indians range themselves territorially with those of Tennessee and also Ohio, in whom both Blake and Russell found a high incidence of these growths; the Kentucky Indians show a slight similarity to those of Tennessee; the Dakotas are unconnected with any earlier or present groups reported on in this connection; our Polynesians harmonize with Welcker’s and Davis’ Marquesans, who also showed a high incidence of ear exostoses, and seemingly with the Polynesians in general. From all the preceding it is plain that the highest frequency of the condition is found, on one hand, among the old aborigines of the North American continent, and on the other hand, in Polynesia. In North America ear exostoses were most frequent in portions of the north- central, south-central and central-eastern parts of the present territory of the United States, in the Columbia basin “ flat-heads ”’ (Oetteking), in some parts, at least, of Mexico (Studley: Coahuila) ; and in South America, in Peru. Peru, which was supposed to head all the Ameri- No. 6 EAR EXOSTOSES—HRDLICKA 23 can groups in the frequency of these growths, is seen to be barely in the middle of the American range of their frequencies. Additional details of importance regarding the development and prevalence of the abnormalities under consideration on the American continent, and of their significance, will be dealt with in succeeding sections. DETAILED DATA AGE Older observations —The youngest subject in whom an exoStosis in the external auditory canal has so far been observed, recorded by Field (1878a), was a girl of 3 years, in whom the bony growth fol- lowed the removal of a polyp. This being by far the earliest age at which the abnormality has been found, the case deserves to be quoted in full; the report reads: M. W., a little girl aged 3, was brought to the hospital on July 25th. Her mother states that she had suffered from a severe attack of measles 12 months previously, and that she had since had an offensive discharge from the left ear. I had a few months since removed a polypus. About a fortnight ago, she noticed a hard substance in the ear, causing the child much uneasiness. When she came to the hospital, a small pedunculated osseous tumour about the shape of a pear was discovered, almost filling up the meatus. The individually recorded cases in the young and up to about the age of puberty, that I was able to find in the literature, are as follows: Field (1878a), in a girl of 3 Brindel (in Sabroux, 1901), in a boy Alexander (1930), in a boy of 9 of 13 Tod (1909), in a boy of 10 Alexander (1930), in a girl of 13 Krakauer (1891), in a girl of 12 Kessel (1889), in a girl of 14 Garrigou-Désarénes (1888), in a boy Ferreri (1904), in a girl of 14 of 12 Bezold (1895), in a boy of 14 Karewski (1892), in a girl of 13 Green (1879), in a boy of 14 It may be worthy of note that of the 12 subjects for whom both the individual ages and sex are given, 6 were males and 6 females, a rela- tion which, as will be seen in the next section, does not hold later in life, when the exostoses are much more numerous in the males. Bezold (1895, p. 48) mentions an isolated case in a boy of 14; and in addition West (1909) reports a case “in a child of 12”, who was probably a girl, for the term “ child” would scarcely be applied to a boy of that age. The great rarity of ear exostoses in children appears most strik- ingly from the data published by Bezold (1885). Examinations by 24. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 aurists for ear troubles in 9,939 school children failed to show even a single case of the tumors. The detailed data were: Ler (el ote a ER R= 1 ee ee 1,055 children, no ear exostoses Wei Stubemant ahi e aps osc s-c.s.a 5,905 children, no ear exostoses BezoldsMininichtesaaeene cee cae. 1,918 children, no ear exostoses SOUGHT Ot Ante orciciniy heads seis 570 children, no ear exostoses INorrellyeWaSwAL 2o.cieeia i.e. 491 children, no ear exostoses Kessel (1889) believed that the growths appeared between the ages of 10 and 13, or later—most frequently about the age of puberty. Erhardt thought they originated during the period of the ossification of the canal; and Edward, of Berlin, thought similarly that these exostoses could originate only in the skeletal parts still in the process of formation—opinions opposed by De Rossi, Ferreri, and others. Braunberger (1896) stated they developed both before and after the ossification period of the external meatus, though they are more common in advancing age. Lake (1898) thought the “ hyperostoses often appeared latish in life.” According to Bezold and Siebenmann (1908, p. 102) “ they develop nearly always after puberty.’’ To which Bezold adds, “ In 170 cases which I observed up to 1&96 there was none below 15 years. Neither did I find a single case in public schools or deaf-mute institutions.” In another place, however, as noted above, he mentioned a case in a boy of 14. Gray (1910) states that the bony growths in the ears are ‘‘ seldom found in children.’’ Alexander (1930) failed to find these growths in children, even in those whose parents were badly affected by ear ‘ exostoses. In contrast to this Marx (1926), who conceives a substantial dis- tinction between hyperostoses and exostoses, regards as an essential mark of hyperostoses that they occur in early childhood, whereas the exostoses develop in the adult. Von Troeltsch’s (1873) cases “‘ were predominantly in the middle-aged”; and Whitney (1886), in his Indian series, found them “as a rule in men past middle life.” A most helpful record in this connection is that of Korner (1904, p. 105), who, although going into no details, gives a list by ages “ of the hyperostosis and exostosis cases ” observed by him in his private practice, “in 1,000 consecutive patients with ear diseases.” The data follow: No. 6 EAR EXOSTOSES—-HRDLICKA ty Cn “ Hyperostoses and Exostoses Cases” Observed by Korner Subjects with exostoses Bee Subjects aad perenne number of growths Number Percent pe MR renee ches onset aus? auiseheaislcunrrev es eytoKe ate) 2 age 81 = © Ga OM Rr ee tectet re Che oi eave cla Oe 113 A= ih MAT UN fas iak way HRN nips BU Geman (alee SANS She 116 6= 4.8 MODOC eee teecre fo esis ote ereea wis es shieke, ct 115 i — Ss 22 rses eiehe it es oe eres SA Ute whee SS 78 ie OR 2O—Z Os alathepe wenera sisjateioe Bear Slave ya « eke Gd eee 719, 17 = 13.6 BUSS Ciccone clem so eonp o60.0bn Moo doo oon 74. 12 = 9.6 BOAO ite eet ot are epcre oe a Secretar I ba 74 1 =] 17.833 ANTE A' Stay tefishntcctereirs svasche, eraashes aueeramiees Sys 68 13 = 10.4 AGS Ours cise sess) Oro amaerneicos se ELTe 61 7= 5.6 Ee Ofer rence oe in se ech ehenncetalce erat 38 8= 6.4 GOO Sis) ie] si ohare as «aks cre eyote) ox sieydlln iy house 45 10= 8.0 OMS OS erstorpigs cnc haart aiiot foarte, oiket ta. 29 Alva) a BD CGS OMe ci Med eres oe aera 19 5= 4.0 CONDE FAO'S oles oe CICA OME eae a loca ae 12 I= 0.8 I ,000 125 = 100.0 To the above I am able to add data which I have culled from addi- tional clinical reports, showing IOI cases where some information as to age was given. Of these, in 25 cases the subjects were merely marked as “adult ”’, or “ of mature age’, or “elderly ”. The 77 sub- jects whose age was stated range as follows: Age of Patients (White) With Ear Exostoses Age Subjects Moree BelOWwalOne shinee hier ccna kote aaa 2 2.6 NO 2O Rare etme oad citie sieinds Stabe aneee 13 16.9 2 BO ae eu Gey Acie cmapes ek alee e: Se sae BIO 19 AT, AeA OM gay atass crandheningekayett « SAL Ue oreo 15 19.5 AD SOME ei ee atten seen ce te ae ole 14 18.2 BOOP eu da smear kcucih eta) ne Kshs Gancad Ginyeto oats 8 10.4 OU OS Fe Peery aioe Lin ie sutlene easel ah onal ous irae 6 Fes) EN DOVELOS MEET AOR Ech rN toe ee The greatest frequencies of the bony growths in the external audi- tory canal are registered, it is seen, between the ages of 20 and 50. Their period of prevalence extends chiefly from 16 to 60. They are rare below to and but one had so far been recorded above 70, though the distribution in our table indicates that a few cases do probably occur above that age. The condition or “ disease ’’, if it can be called such, is therefore, it is clear, dominantly one of the earlier to middle adult life, and has 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 very little if any connection with senility and its disorders—another point of importance. It should be borne in mind, moreover, that the clinical age reports embodied in the above data are the ages at the time the subjects applied for treatment, and that in all the cases the bony growths in the ear began earlier. The data therefore show the ages at which the tumors reached development that caused the patient to apply for relief. Should we wish to learn the age at which the ear exostoses began, it would be necessary to shift the whole scale of ages in the direction of less years. How much to shift, on the average, cannot be estimated. In individual cases ear exostoses were known to the patients to have existed for many years before they gave sufficient trouble to lead the subject to the aural surgeon. The matter can only be settled, it would seem, by prolonged future attention to this point. New observations —Excluding the Eskimo, the specimens in which both meatus cannot be examined, and the very young, there are in the National Museum 335 skulls of Indian children between the ages when the tympanic portion is fully added to the bony ear and when there occurs the eruption of a permanent M,—in other words between about 2 and 12 years of age. The material, like that of the adults, is mainly pre-Columbian, and a good proportion of the skulls show one or another of the three kinds of deformation. A careful scrutiny of these specimens showed 334 without any trace of ear exostoses. The sole specimen showing the abnormality was a pronounced “ flat- head” from Santo Domingo, with lateral permanent upper incisors and permanent upper molars about to erupt—hence about 8 to 9 years of age. In the left meatus of this skull is seen on the posterior wall, tympanic part, a small but distinct abnormal nodule, which in all probability would have later grown to a definite exostosis. It may thus be concluded that, among the old American Indians at least, tympanic exostoses did not develop, or did so but exceedingly rarely, in childhood. Even in the Indian adolescent of our collections, however, these growths are not very evident. They belong essentially to the earlier half or two-thirds of the adult life, though they not seldom begin before the adult stage is reached. Their maximum frequency and development in the Indian belong principally to the ages of about 30 to 60. They are infrequent in older subjects and, as in Whites, it is plain that they have little if any relation to senility and its path- ology. All of these facts are of etiological importance. No. 6 EAR EXOSTOSES—HRDLICKA 27 SEX Since Bonnafont (1868), all writers who have dealt with the sub- ject agree that ear exostoses are considerably more common in men than in women. Thus Delstanche (1878, p. 14) says that they are “much more frequent in males,” and practically the same words are used by Von Troeltsch (1881, p. 141). Whitney (1886, p. 442) found “the subjects were as a rule men’’; for Kessel (1889) the growths were “ decidedly more frequent in the males.”” Schlomka (1891, p. 16) and Sabroux (1901) express themselves exactly as do Delstanche and Von Troeltsch. For Braunberger (1896) it is “certain they are more common in males.” There are many other expressions to the same effect. More definite data on the subject, in living Europeans, are given by Korner and Bezold. The latter says (1908, p. 102): “‘ The male sex is affected much more frequently than the female. My statistics show a proportion of 11 to 1, those of Korner of 3.5 to I.” The individual clinical reports are frequently negligent as to sex, but from those in which sex is stated, the indications are very conclusive. Out of 89 instances that I was able to gather in which the sex was given, 70 of the individuals, or nearly four-fifths (78.7 percent), were males, and but 19, or a little over one-fifth (21.3 percent), were females. In the old craniological observations on non-European peoples sexing has been attempted in but a very few cases and that inade- quately ; but as far as the data go they show among those affected with ear exostoses a decided predominance of males. New observations.—The new materials reported upon in this work have all been sexed on the basis of ample experience, with all possible care, and often with the aid of the rest of the skeleton, so that the records may safely be held reliable to within less than 5 percent of possible error. The results are given in the table on page 28. The data show that: In all the groups, Egyptian, American, and Polynesian, ear exostoses are more common in the males than in the females. The total frequency of the growths in a group appears to have little if any influence on their relative frequency in the two sexes. The proportion of occurrence in the two sexes differs considerably between some of the groups. Much of this difference is probably con- nected with the unequal and not always adequate number of specimens, but apparently there are also other reasons, the nature of which re- mains obscure. In the two Polynesian groups the relative frequency of the growths in the females is higher than that in any of the American series. 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 The clinical records cannot be directly compared with our figures ; they lack one of the essentials, which is the proportion in each sex of those with ear tumors to those without ; but as the proportion of the two sexes in the White population at large is much the same, the rela- tive values of the cases reported are probably nearly correct. They Relative Frequency of Ear Exostoses in the Two Sexes Total P f skull Group dott paiet Gan eto fae Male te (in order of frequency unt cb of skulls cma f t ) of skulls ith (male awaears examined exostones Males Females = 100) Hoy ptlallernct rer cone are - 454 9 Bar ews 272 American: BL SNGHMON te eee. hee ce eters I ,000 2 0.8 cae cere OldgPRuchlowne veosans ssa 500 12 2.8 2.0 Fplod INES akotay carte serosa hehe 29 2 1b ie Toe oe loniGama serie ae aie si: f 395 35 1367, Sieis} 24.1 CaltGinians yahoo 3h ato asa s 435 46 15.2 Sa 7/ 37.5 INI States) saeror vey: cnc oe okeccie II2 13 16.0 10.2 63.8 IG Hheng ciaig eats Sigal. Ck oe Cn 3,651 522 222 6.3 28.4 WEES TaN USSR pict Ga ene ee 65 14 28.6 1g). 46.5 WOUISIAN Ameya tie os ces hes 61 15 46.4 6.1 ie ya PADICATISAS atest Me Meee varnii css o8 173 47 38.6 16.7 A338 SsDakotavallis.ic2ee. 6 012 109 30 38.2 Ws 9/ 43-7 S. Dakota, Mowbridge...... 76 23 Atins 21.4 51.8 PREM EMEKV Ae sents ak eke oe rere) 29 48.0 m2 5 26.0 Polynesian: Mew Zealand ty. see : 19 4 25.0 18.2 72.8 awa ees Actes cokes 148 39 28.4 24.7 87.0 indicate that for every 100 White males with ear exostoses there are about 27 females with that affection. In most of the Indian groups and in both the Polynesian groups the proportion of females was decidedly higher, a fact for which there must be reasons as yet undetermined. FREQUENCY OF EXOSTOSES IN THE TWO EARS Our material permits us to show definitely the number of individual ears affected in the different groups, and the proportional involve- ment of the two ears in the affected skulls. The next table gives for our series of groups the proportions of affected ears in relation to that of the affected skulls or individuals. It will be seen that, barring a few exceptions, in groups where the frequency of individuals or skulls with ear exostoses is smaller, the percentage of single ears free of the growths is larger than in those groups in which the proportion of individuals or skulls affected by No. 6 EAR EXOSTOSES—HRDLICKA 29 these abnormalities is larger. In other words, in general, the indi- viduals or skulls that are more subject to ear exostoses will show not only absolutely but also relatively a larger proportion of ears involved than the groups where the growths are less common. Were some of our smaller series as large as are others, the figures would probably be more regular and even more convincing. The meaning of the phe- nomenon can only be that there must exist something like an oto- exostotic “inclination”, which differs in quantity in different racial, geographical, or other groups; and that the larger the quantity of this + in a group, not only the more individuals, but also the larger percentage of individual ears, will become subject to the growths. The affection acts thus as an entity—perhaps more accurately a “ dia- thesis ’’,” a conception which will be found further strengthened when we come to consider the numbers, masses, and character of the ab- normalities under discussion. Relative Proportion of Skulls and Ears Affected by Ear Exostoses in the Different Groups Percentage Percentage Percentage : Group Skulls of skulls Ears of ears of ears in order of frequenc F F : : affected vs. : of ear exostoses) z aeeee pulbeay. ee Lh hs percentage OM RIAN tis utara katte Heh 454 2.0 908 tog 66.3 American: ESKIMO Pei ne ae I ,000 On2 2,000 0.2 (100) OLGEPiIe DOR siete: ee «ie ieee. 500 2.4 I ,000 1.9 79.2 I os) 0 5G 8 Sie is eeepc ac 29 6.9 58 3.45 50.0 ilOnidarpepresiatic toh ey etc se avele 395 8.9 790 6.7 75.3 ASAE ceo ees aces Gra. oe ssl ens 435 10.6 870 7.9 74.5 ONES otates! i. seine ea ans 112 1t.6 224 8.5 TBs Renee sen as atten dieit We 3,651 14.3 7,302 Tt ai7/ 81.8 BU SMMIN AG a cote ice scan ie 6 eek 65 21.5 130 19.2 89.3 PROS AA re ew he oe eae eas. 6 61 24.6 122 28 86.6 PATIKATISAS rata Torsrnare trotted 173 Dae 346 2147 79.8 SaDakotaralli@s ces ros 109 27.5 218 22.0 80.0 S. Dakota, Mowbridge...... 76 30.3 152 24.3 80.2 USGI CCl 9. cas cae ea ae 90 22.2 180 28.9 89.8 Polynesian: Mew Zealand: 553023. ae0's cick 19 Aisi 38 18.4 87.2 DU Wallles shoe ace Paik ae coteuecntea ees 148 26.4 296 20.3 76.9 SIDE Since the early history of ear exostoses, it has been known that such growths have a tendency to occur bilaterally, but opinions differed as to the frequency of such an occurrence. “In the sense of “‘a constitutional predisposition or aptitude for some particu- lar development” (Webster’s New Int. Dict.). 3 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Delstanche (1878, p. 10), stated that though the exostoses showed a tendency to occur on both sides, in his personal experience uni- lateral cases were “much more frequent’. Schlomka (1891, p. 16) and Braunberger (1896) have also reached the conclusion that they occurred more frequently unilaterally than bilaterally. On the other hand, Von Troeltsch (1881, p. 141) saw them to be “ predominantly bilateral”; Sabroux (1901, p. 35) says they were “ very frequently bilateral”; and there are other statements of similar import. Blake (1880) gives the first exact data on the subject. Of 231 old Tennessee and California Indian skulls, 38 had ear exostoses: in 14 (36.8 percent) these were bilateral ; in 9 (23.7 percent) in the right ear only; and in 15 (39.5 percent) in the left ear only. Attempts at statistical estimates in Whites were made by Korner (1894), who found the exostoses in 64 percent of cases bilaterally, in 36 percent unilaterally ; and by Bezold (1895, p. 48) who found them in 54.4 percent of cases on both sides and in 45.6 percent on one side only. Bezold observed: that occasionally, in cases originally unilateral, in time a development of the abnormality would also take place in the second ear. As to the side in the unilateral cases, Ferreri (1904) thought the growths were more frequent on the left. New observations —The table that follows gives the involvement of the ears as to side in our material. Side Percentage of skulls Groups Percentage with ear exostoses of ear exostoses) exostoses | Bijateral Right ear | Left ear only only Space ciety eile occ ett 454 2.0 23.23 2252 44.5 American: 12S iTNVals een ots onde, ceo eR eo I ,000 0.2 (100) ican Beat @ldbRueblomorosmen. ego 500 Bish 58.3 16.7 25.0 NeDakotare sas situs faxes 29 6.9 (Num|ber inade|quate) Hlonidanerraraecocepe oe ot ee 395 8.9 51.4 28.6 20.0 Califouniabaer ater: see 2 435 10.6 50.0 23.9 26.1 NIB States ert sites Sieve 112 Wy asy 46.2 15.4 38.5 PEK ate to aA ieee a sesne es 28050 sulla ss 63.6 16.1 20.3 Wing intaewrs eins rary eet: oh 65 215 78.6 72 14.3 ouisianare meer eee. er 61 24.6 73S nes 13-3 Arkansas@ econ ae sae. 3.3 173 Dae 63.8 19.2 17.0 S) Dakotaralle en cewretee sc 109 27S 60.0 20.0 20.0 S. Dakota, Mowbridge ...... 76 30.3 60.9 Gp adel 217, IWentucky ae. pamece eo eae go 2202 79.3 13.8 6.9 Polynesian: New Zealandin. ssn sedan: 19 Zee 75.0 ies 25.0 ELE Wall na ees Beha e ae cae weal 148 26.4 53.8 15.4 30.8 No. 6 EAR EXOSTOSES—HRDLICKA 31 The data on page 30 show the following conditions: I. In general, in a group the larger the number of individuals af- fected by ear exostoses, the more commonly both ears are involved. 2. The frequency of bilateral as compared to unilateral involvement varies from 33 to 79 percent in the different series of our material. 3. When but one side is involved, there is a perceptibly greater tendency for the abnormalities to develop in the left ear. In the 14 groups of our crania, in 9 the involvement was more frequent on the left side, in 3 on the right side, and in 2 groups it was equal on the two sides. The proportions of bilateral involvement in our series agree fairly closely with those of Bezold (54.4 percent) and Korner (64 percent) on European Whites. Through all this the affliction again impresses one as something of a pathological entity. As it is probable that some proportion, at least, of the individuals whose skulls enter into these series, if they had lived longer, would have developed the condition bilaterally, it may be assumed that the general tendency for the tympanic exostoses is to involve both auditory canals; but they evidently tend to start more frequently on the left side. Why this should be so is not apparent, but it is probably con- nected with nervous and vascular factors. The very apparent tendency toward a bilateral involvement should be an important factor in the study of the deeper causes of these formations. NUMBER Ear exostoses may be single or multiple. Delstanche (1878) en- countered up to three in one ear. Von Troeltsch saw “ frequently several in one ear, arising from different parts of the wall.” Sabroux (1901, p. 35), from the data he gathered, believed them to be “ ordi- narily single’, though there might be two or even three in one ear. For Kerrison (1913, p. 125) they were “ multiple oftener than single.” Blake (1880) and Korner (1894, p. 107) alone give some statistics on the point. In Blake’s 39 cases of ear exostoses in Tennessee and California Indians, 54 of the 78 ears showed the growths: in 38 ears (70.4 percent) they were single, in 12 (22.2 percent) double, in 3 (5.6 percent) triple, and in 1 (1.8 percent) multiple. In 41 ears Korner found the exostoses 32 times (78 percent) single, 8 times (19.5 percent) double and once (2.7 percent) triple. The two records are remarkably harmonious. Aside from Blake’s there are no data on this subject in the published reports of a more anthropological nature. Among the clinical records 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 in the literature I was able to find 84 cases} or rather ears, in which the number of exostoses was given. The results follow: Number of Exostoses in Individual Ears of White Subjects Recorded in Otological Literature Number of records Exostoses on individual = a ; — ears Single Double Triple or more 84 44 27 13 52.470 32.1% 15.5% It seems plain from the above data, small in number and imperfect as they are, that among the Whites, in over one-half to two-thirds of the ears affected by exostoses in the external meatus which come to the attention of the aural surgeons, there is but one of these growths, and in the remaining number there are two or over. New observations——A much more satisfactory record can be pre- sented in connection with the new observations that are given in this work, though on different racial groups: Number of Exostoses in Individual Ears Affected with Such Growths Skulls Percentage of exostoses in individual ears Group Skulls with ear exostoses 1 2 3 Over 3 IDEAWDISENNG on oro Ge ooo enc 454 9 83.3 8.3 ee 8.3 American: eS kai OP Meus sects Gees I ,000 2 (100) ae OldgBueblozse.ssn4-- 500 12 100 : Bie oe INEGIDalkotalere aise ac 29 2 (100) ahs Re ae A lonidamine emma 395 35 la 24.4 2.0 .O Cahifonniaeeecescec. - 435 46 737, 175 7.5 Wo INEsiStatesiss sin. soo 112 13 W277, 26.3 tare ; Renee ee ean tickers 3,651 522 46.3 38.6 14.6 0.4 Winging aie) ss 3 sic * 65 14 52.0 36.0 4.0 8.0 ICOUISIananys ane ee ee 61 15 732 26.8 a pW aps Akan Sasaiv cise cin aise 173 47 Aang 25.3 1.4 ae SyDakotaralleeecin: 109 30 75.0 16.7 Le 8.3 S. Dakota, Mowbridge 76 23 81.1 16.2 ane Dey INGnIGHeKyao are scres go 29 ae 26.9 15.4 ae Polynesian: New Zealand........ 19 4 (42.9)) | (14:3) Gag) Gases FlAaSvallieeanyerorereste sens 148 39 56.9 36.7 4.9 tek It will be seen that in all the larger groups a single outgrowth in a meatus is the most frequent condition. In our Pueblos, curiously, it is the universal, and in our old Egyptians a nearly universal, condi- tion. But in most of the other groups two exostoses per meatus are also fairly frequent; three are scarce, and more than three are decided rarities. No. 6 EAR EXOSTOSES—HRDLICKA 6) The single tumefaction-exostosis is, as a rule, the initial stage, but a second enlargement, generally from the opposite side of the tympanic portion, will soon follow, the third and perhaps additional growths developing later. In no less than 6 of the 12 Indian series the percentage of single tumors is practically identical. In three of the remaining groups, Peru, Virginia, and Kentucky, and in both Polynesian groups, the proportion of cases with two or more growths is exceptionally high. The Peru and the Kentucky material especially shows many ears with three exostoses. Two notes of caution are here necessary: 1, In some specimens with one ear exostosis, had the subject survived to older age there might have taken place a development of additional growths; and 2, where the canal is nearly filled by one, two, or three exostoses, other growths, smaller and more posterior, cannot be seen and thus escape enumeration. This means that the proportions of cases with more than one exostosis would be somewhat higher were we dealing with those who lived their full span of life; and that the proportions of two and especially those of three or more growths are in reality in some measure higher than those given in the records; but the excesses in either case could hardly be very material. The chief meaning the above results convey is once more that the ear affection under consideration appears in the form of an entity or “ diathesis ”, which manifests itself practically identically in a large proportion of different geographic or tribal groups of the same race, but which may also, through some peculiar causes, differ substantially in other groups of the same people. LOCATION As in other respects, so in this also there are in the history of ear exostoses some variants of experience and hence of opinion. According to Toynbee (1860), ear exostoses may develop in any part of the length of the tube; but the part from which they most commonly originate is about the middle third of the passage. .... Occasionally the posterior wall affords the point of origin to the tumour, and then it not unfrequently resembles a simple bulging of the wall. In other cases, a similar tumour is also developed from the anterior part, and the two protuber- ances meet and lie in contact in the middle, leaving an inferior and superior triangular space at the place of the original opening of the tube..... The tumour may grow from the upper surface of the tube..... Two or more tumours, again, extend sometimes from various parts of the circumference of the meatus, and converging towards the center, fill up nearly the whole cavity. 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Delstanche (1878) says that the growths may develop from any point of the meatus, but arise most frequently posteriorly. For Blake (1880), most of them are on the posterior wall; for Ayres (1881), they are in any portion of the canal but most frequent posteriorly ; Von Troeltsch (1881, p. 141) states they “ occur as well at the be- ginning of the bony canal as close to the drum . . . . by far the most often from the posterior and superior, less often from the anterior wall.” R. Virchow, and after him Hartmann, Ostmann, Korner, and Marx, held that the abnormal bony growths in the external auditory canal originated generally (or even invariably) from the borders of the tympanic bone, where they lay on and fused with the squamous portion of the temporal; some of these authors, however, had observed also occasional exostoses in other parts of the canal. Moos (cited by Gold- stein, 1898), in his cases, observed that they arise “ invariably from the upper wall of the canal, almost at its outer end’”’; in Jacquemart’s (1889) practice they were ‘“ encountered most commonly at the in- ferior part of the meatus.” Among additional statements, some of which merely repeat others, the following may be quoted. Schlomka (1891, p. 16): “As to their origin, these exostoses arise most often from the postero-superior wall of the auditory canal . ... more seldom from the anterior, most rarely from the lower portion of the same.” Sabroux (1901, p. 34) says that they develop anteriorly and posteriorly as well as superiorly and inferiorly, “ but it is at the union of the cartilaginous with the bony part (of the meatus), or again quite close to the drum, that we observe them most often.” Ferreri (1904): ‘‘ Most frequent pos- teriorly and superiorly.” Kessel (1889, p. 285): “‘As far as the place of origin is concerned, the globular exostoses arise from the posterior and superior wall, the cylindrical ones extend along the floor of the meatus parallel to its axis, while the osteophytes occur preferably on the superior part of the inner portion of the canal.’’ Kerrison (1913, p. 125): “ Oftenest on the anterior and posterior walls. ... . When one growth only is present, the anterior wall near the attachment of the drum membrane, is the most common site.” Dahlstrom (1923, p. 213): “Most of the exostoses were found approximately on the boundary between the cartilaginous and the bony parts.” Burton (1927): ‘“ By far the larger number of aural exostoses are found along the suture lines between the canal and the rest of the temporal bone.” Oetteking (1930, p. 249): “The pea-shaped restricted to marginal area of meatus; more or less pedunculated anterior or posterior; the crest-shaped is elongated and extends from without inward.” No. 6 EAR EXOSTOSES—HRDLICKA 3 cn For Moller-Holst (1932, p. 96), who had the advantage of observ- ing numerous cases in skulls, “ the exostoses of the auditory canal occur in the outer, middle, and inner portions of the canal. They are located on the tympanic bone in different places, mostly on the boun- dary of this bone and the squama or pars mastoidea. They seldom proceed from the squama.” In his material, “they occurred with about equal frequency anteriorly and posteriorly in the tympanic part; one-third of the cases showed them anteriorly, one-third posteriorly, and the last third both anteriorly and posteriorly.” Older records.—The only actual record on the location of ear ex- ostoses within the meatus is that of Blake (1880). In 45 Indian skulls with such growths, from Tennessee, California, and Arkansas, out of a total of 82 exostoses 55 (66.3 percent) were posterior, 24 (28.9 percent) anterior, 3 (3.6 percent) antero-superior, and I (1.2 percent ) postero-inferior. New observations —The location of the exostoses was noted in all of our material. It was as shown in the table on page 36. Notwithstanding some significant differences, there is a remarkable basic agreement in the figures, and also between that part of them that embraces the bulk of the American Indian material and those of Blake on the same race. It may now definitely be said that ear exostoses in the American Indian (with a few exceptions), regardless of the amount of total involvement by the growths, will be in something over 60 percent (of all the exostoses) posterior, in from 20 to 40 percent anterior to antero-superior, and in a small percentage—and that in but some of the groups—postero-inferior or antero-inferior. Superior (squamous part) outgrowths occurred in but three of the series, inferior out- growths in but one and that only in a single instance. Some observers would perhaps be inclined to class some of the cases included in “ posterior ”’ as postero-inferior or postero-superior, but that would not alter the main significance of the above showing. This is that, in general, in the American Indian, approximately two- thirds of the ear exostoses arose from the posterior end or portion of what was originally the tympanic ring and later developed into the tympanic bone; that nearly one-third of the growths arose from the anterior end of the ring and subsequent bone; and that in but a few groups was there apparently any tendency toward a development of bony excrescences from the squamous portion of the meatus—though the incidence and proportions as given in the table for this item would probably be augmented somewhat if an examination of the whole squamous part were always possible; and that the floor of the meatus VOL. 93 g€z 0°61 $-zS 6¢ gti auiss susie ite ecole) Si0) aah a EMPEY S L-9z 0°02 Cres t 61 "so 57 +s pueleaz Man © :uRISIUA[Og & a ria L0z 6°S1 z‘z9 6z OG a's ule eae ce Ayonquay =) ah Z v9 6° V1 G'bZ tz gL aspluqmoy ‘ejOxeq ‘S is Seen 8°8 6°z 6S OFL1 Ca £19 of OO Sie se ‘Te ‘eJ0XxeGq °*S [S) sai o. l'z 9 tr 9g SI I'z 9 ‘Sg LY Ce ee |e eee et svsueyIy n aun Gulic 2) mea £9 6a C1 19 su Cue visceral ace e BUPISINO’] = gt €-pr g€z oon 1°ZS TI S9 salto eve Meals Magione PIUIBII A, es] 10 € Sz 6°of Ovl Lev ces LS OBC a tape cma ae ods niaq vA one ud GELS cz CT FA\ Wifes ell (poe aoeO $9}P1S "IN < o:Sz bL yt 0°99 ot CO ie || SEU Cee PIUIOJI[E-) a € vz o/s va 6% £°¥9 Se. COCg goat cbse Se le eployy Oo eiagie sais eyreics (001) Z 6z ele) eftcthestnl ysitane lr P}0AeC "Ni 4 OO1 zI Gosia) Sy pee ees olgeng PIO = Ool z 000‘ Te oe Remade iain Sela nAonion fos Owlysy = PURILIOWYy < z 9° Text etn a) farne V . rae) 6 Sr ie) 4, 0.0/6 6) 1040.6 810 U6 (ee ueiqdA3sy [e) ——— i 10lajuy iowedns Esau eae IOloyUy ce ee 101194S0qg Sates Wigaliaide = s[[nys Jo s[[nys jo ees) a SYJMOIZ 9Y} JO 1oquINU [2307 JO 9deyUVI19d UT ‘SasO}SOX9 JO UOIQRIOT te Ua eS UUN SNIDIPY IY] ULYJIM Sasojsoxy AD ayz fo urs1U¢C fo avj1q 36 NO. 6 EAR EXOSTOSES—HRDLICKA oF is practically never primarily involved by these abnormalities, though it may be secondarily involved when the posterior or anterior exostoses are large and diffuse. In the old Egyptians of 4,000 years ago there is the same predomi- nance of posterior growths, with a smaller proportion of anterior, and no other. In the Polynesians conditions differ but slightly from. those in the Americans: there are somewhat less of posterior and correspondingly more of anterior and antero-superior exostoses. The data demonstrate once more that, although there are some group differences in the location of ear exostoses, this location discloses certain general tendencies, which in their turn support the conception of the process of these growths as a definite and separate abnormal entity. Observation along this line on large numbers of specimens leaves a strong impression of basic definiteness and rule, both as to location and mode of development, which is radically opposed to any incidental or purely local origin of these formations. SIZE In size the ear exostoses vary, according to Wyman (1874) from “a pin’s head to that of the whole caliber of the canal”; according to Whitney (1886), “from a simple thickening all stages can be traced, up to the formation of round osseous growths as large as pease, which in one or two cases have completely blocked up the opening ’’; according to Russell (1900), from ‘“ minute nodule to large tumorous growths”. Sabroux (1901) says that their volume varies much—from a simple protrusion on the walls of the canal to a complete obliteration of the same. For Gray (1910) their size ranges from “a pinhead to that of a bean”’. Dahlstrom (1923) states that they “‘ seldom reach such a size that they would completely close the meatus.’ Moller-Holst (1932) says that “the size varies from that of a pinhead to exostoses that involve nearly one whole wall of the meatus; while in height they range from very low to those that project up to 7 mm.” There are additional statements on the point, but they merely repeat more or less what has been given here. New observations —From our observations, unimpeded by any soft parts or wax or discharges, it is possible to state as follows: The growths differ from small distinct abnormal bulgings or little circum- scribed exostoses to bony tumors that almost occlude the meatus, or even protrude beyond it. In no instance has an absolute occlusion of the auditory canal been seen, but when in life the parts were covered 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 with skin the closure in some of the cases must have been about com- plete. Large growths are frequently accompanied by smaller ones. The circumscribed exostoses never coalesce, and they never become fused with the wall they reach in their growth. An attempt was made to record the exostoses found as to size. Such a procedure is difficult and calls not only for extensive experience but also for a clear notion as to the range of variation of the growths. With these conditions satisfied, it is possible to arrive at a rational classification which, although incapable of mathematical accuracy, is nevertheless really useful for conveying a fair picture of the condi- tions. The whole range in size of the growths was divided into small, moderate to medium, and large. The ‘“‘ small ”’ ranged from that which could be distinctly diagnosed as a new growth to those grades where the formation began to be too much developed to be still readily included in that class; the “ moderate to medium” were those from the upper boundary of the “small” to such as would occlude up to a half of the canal; and the “large’’ were those growths that oc- cluded more than half of the meatus. The results obtained are not without interest : Size of the Ear Exostoses sa Percentage Size of the bony US Group of skulls Ot sulle 7 examined exostoses Small pias toe Large p Percent Percent Percent Bowpblaitns ete cre ssa hs oon 454 2.0 76.5 23.5 American: Skimpy ake ace ces oO oer I ,000 0.2 25.0 75.0 oat OldePueblovicaeugc. 225 ae ees 500 Bol 100.0 aha is IN-eDakotaiaccss. 3.19. wea woe 29 6.9 50.0 50.0 fe Ploridaemceie cece snakes ak a 395 8.9 67.2 25.7 Gol Galittionmitae sg. :toio ss fee ake s 435 10.6 78.8 18.1 B2 INE Stateses en f . gama oi T12 11.6 75.0 25.0 acne IBELU cen creer s aris vis dale eta: 3,651 TAs 50.9 Aas 6.6 WIRGIMID a teeta tracy cin oaine 65 215 52.4 40.5 Fal IEOUMISIANA yey fee scares cee 61 24.6 78.1 18.7 301 INT Kan sa smear prer rie Giiciee es ae 173 272 65.7 28.1 6.2 SyDakotayall i 5 eo. aes 109 BG 60.3 39.7 Lat S. Dakota, Mowbridge ...... 76 30.3 55.1 44.9 xi Wwentuckyearraner actin caciae go Bo 45.1 50.0 4.9 Polynesian: INewsZealardreri nares och ast 19 PEI 86.7 133 Flea Wi ttre te elise nstene cere.) ae ts 148 26.4 96.7 Bh There are some very perceptible and significant differences in the above figures. In the old Egyptians, the Pueblos, and the two Poly- NO. 6 EAR EXOSTOSES—HRDLICKA 39 nesian groups, in the two Dakotas and in the northeastern American States, there were no exostoses that could be recorded as “ large ”’. In the Egyptians, the Pueblos, and the Hawaiians, in particular, not- withstanding their widely differing total involvement by the growths, the exostoses were all or almost all small. The greatest collective “massiness”’ of the abnormal formations occurred in Kentucky, Virginia, Arkansas, Florida, and Peru—again under widely differing total involvement. The causes of the above differences are not clear. If it were prac- ticable to divide the basic causes of these exostoses into hereditary and those checking, or again favoring or exciting, their development and growth in the predisposed individual—as may be correct—then it could be conceived that the differences in their total bulk in a group were due proportionately to the checking or favoring factors. Another influence that enters into the subject is that of age. The growths need time for their development to a large size. Of two groups equally subject to the exostoses the younger would presumably show less mass of the growths. It is probable that the mean age of the affected skulls is not the same in all the groups. But age alone could not possibly account for all the differences encountered. One item is clear from the results shown in the preceding table, and that is that the numerical involvement by ear exostoses in differ- ent groups does not always go hand in hand with the collective massi- ness of the growths. SHAPES Ear exostoses occur in many shapes, and these merge with each other so much that any definite classification appears impossible. Blake (1880) believed that he could recognize two main forms, the “rounded” and the “ flattened ”. Von Troeltsch (1881) divided the better differentiated ones into broad-based and pedunculated. Kessel (1889, p. 285) states they are “ multiform ”’, but fall into three classes, namely, hyperostoses, periostoses, and exostoses; under the latter “are understood more or less circumscribed tumors that grow espe- cially in one direction and stand either vertically to or parallel with the axis of the canal. One can observe rounded, cylindrical, and conical exostoses ’’—but there are also transitional and peculiar forms; there are, too, spinelike growths which have been called “‘ osteo- phytes ”. For Schlomka (1891, p. 16), who doubtless in part follows Kessel, “ one can speak of rounded, cylindrical, or conical exostoses ; but there are many combinations of these forms”, and some of the growths may assume the form of ossified polyps. Sabroux (1go1, 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 p. 35) says that their implantation is various, ranging from large diffuse to one or even more than one “ pedicules”’. Korner (1904, p. 106) believed that “ the globular growths tended to be located more outwardly than the small buttonlike exostoses, to arise from the pos- tero-superior wall of the canal, to be larger than the other excres- cences—pea to cherry size—and to lead readily to the complete clos- ure of the meatus.” Moller-Holst (1932, p. 96) says: “As to the shape, one can distinguish flat, broadly based, semiglobular, globular, and pedunculated exostoses. All the forms connect through inter- mediary grades.” New observations—In our examinations it was found that the ear exostoses show a variety of forms which merge into each other, present no distinct separate classes, and can only be subdivided roughly or arbitrarily. They range on one hand from what can clearly be recognized as an abnormal localized tumefaction or bulge on the pos- terior or anterior tympanic wall to large irregular massy growths with adventitious smaller elevations; from slight oblong ridges directed along the axis of the canal to redundant more or less conflowing welts ; from a recognizable swelling of the postero-superior or antero- superior terminal part of the tympanic ring or bone, to masses that almost occlude the auditory canal; and from little pearllike exostoses to more or less buttonlike or pedunculated and irregular tumors, some of which nearly occlude and some even protrude from the meatus. On the other hand, there were observed no slender spines, or sharp osteophytic growths, and no forms that could properly be called “ cylindrical ”. SYMMETRY There are repeated statements in the literature on ear exostoses that these growths in some cases occur symmetrically in the two ears of the same individual. Thus Dalby (1876) states that “not infrequently both meatus are affected similarly, not only in respect to the presence of these tumors, but also as to their size and number ” ; and they may also “increase synchronously” in the two canals. For Delstanche (1878, p. 10) “ most often, in case the tumors are bilateral, their point of insertion corresponds exactly in the two canals, and they possess also a certain analogy of form and even of size.”” Ayres (1881) says, “ There is seemingly a good deal of uniformity in their development ” ; and similarly Kessel (1889, p. 286) states, “as to the bilateral cases, it may be remarked that they occasionally appear in the same form and in the same locations and therefore symmetrically’; and for Schlomka (1891, p. 16) the bilateral outgrowths “are occasionally No. 6 EAR EXOSTOSES—HRDLICKA 41 entirely symmetric, both as to form and location.” Noquet (1899) in one case saw perfect symmetry on the two sides—the same aspect, volume, point of origin; Urbantschitsch had one case with two ex- ostoses in each meatus, exactly alike. Sabroux alone (IgoI, p. 35) considers them as “ very rarely implanted symmetrically in the two auditory canals.” For Korner (1904, p. 107) they “develop mostly symmetrically ’’; and also for Moller-Holst (1932, p. 96), “they in part develop very symmetrically.” New observations——wNo statistical data have been attempted in this connection, for it was learned that a tendency toward symmetry, where both external auditory canals are affected by exostoses, is the general rule. The likeness extends most to the location of the growths, less to their number and size, and least to the details of their shape. Even where the growths must be recorded as unilateral, there is frequently a trace of a commencing tumefaction in the corresponding locality of the other meatus. This general tendency of the growths toward symmetry on the two sides connects them clearly, in the view of the writer, with the central neuro-vascular system and is another element of importance in the etiology of these formations. RATE OF GROWTH An inquiry into this subject is possible only to the clinician. There are a number of notes on the subject in otological literature. Toynbee (1860) learned that the development of these tumors may frequently be very gradual and “ unattended by any symptoms calcu- lated to attract the attention of the patient.’ Dalby (1876) found that ear exostoses “ may remain without perceptible change for many years.” In the observation of Von Troeltsch (1881), their growth was gen- erally very slow; Braunberger (1896) stated that it was “slow and painless.” Kessel (1889, p. 288) noted that, although usually slow, their growth at times was rather rapid. “It was most accelerated toward the time of puberty; older age retards growth. The growth is sometimes uneven; it can cease for a time and then recommence.”’ In Stewart’s (1901) experience, although such an occurrence was “extremely rare’’, yet on occasion these exostoses could take on a rapid growth suddenly. A summary of all the clinical observations on the rate of growth of ear exostoses is that their early stages are slow, symptomless, and usually ignored even by the subject; that many apparently stop all growth after reaching a smaller or larger size; but that under exciting 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 conditions, evident or not, there may set in a relatively rapid enlarge- ment, which soon is attended by distressing symptoms that bring the subject to the aural surgeon. An interesting further fact is that after an operation and removal of an ear exostosis, in a majority of cases the subject remains free from the growth, but in some cases there is a recurrence of the bony tumor. In the absence of both infection and malignancy the latter event implies, it would seem, a special neuro-vascular influence. SUBDIVISION Since Toynbee (1860), repeated but not very successful efforts have been made at a classification of ear exostoses, either on the basis of causation, of the form of the tumors, or of their structure. The morphological classification alone needs to be dealt with in this section. It begins with Cassells (1877), who divides the growths into two categories, namely, the hyperostoses and the exostoses, which he believes to be separate abnormalities and of totally different origin. This clinically convenient subdivision, notwithstanding voices to the contrary and its demonstrable basic inaccuracy, has become generalized in otology and will probably stay there. Von Troeltsch (1881, p. 144) states that hyperostoses are not clearly separable from exostoses, yet he uses the two terms, applying the first to “a generalized hyperplasia of the walls ” of the meatus. Virchow, though in general admitting the subdivision, appears at times uncertain; he speaks (1889) of a “ hy- perostosis ” in a Tanimbar skull which closely resembles the “ ex- ostoses”’ of the Peruvians. Kessel (1889) follows Cassells, but adds two more forms and gives the following somewhat arbitrary definitions: Hyperostoses, involv- ing the whole extent of the canal; exostoses, more or less circum- scribed growths ; periostoses, ringlike bony growths ; and osteophytes, spinous growths. Pritchard (1891) attempts to divide ear exostoses into: 1, multiple, uniformly smooth and rounded ; 2, multiple, irregular in shape ; and 3, single, polypoid. Hartmann (1896, p. 44) has this to say: Now as it happens that of late years both hyperostoses and exostoses have been described indifferently under the title of exostoses, although they are plainly two different processes; the descriptions of cases vary a great deal, depending on whether the author has had his mind more intent on exostosis or on hyperostosis. That there is an essential difference between the two, is also supported by Virchow, who, with his enormous experience with exostoses, remarked, during the discussion of a paper which I had read in the Berlin Medi- cal Society: “I agree with the reader, particularly in one point which made No. 6 EAR EXOSTOSES—HRDLICKA 43 itself plain to me many years ago during my ethnological investigations of skulls from the northern portions of the Pacific coasts. I refer to the more diffuse hyperostosis which he has exhibited in his first case. That, I should not call an exostosis at all. Diffuse hyperostosis is quite different from a genuine exostosis. Inasmuch as exostoses have a very typical situation, this interpreta- tion is very plain.” Hyperostosis of the external auditory meatus always limits itself to the pars tympanica, whilst the adjacent squamous portion of the bone has no bony pro- tuberance..... . ; The entire pars tympanica may become hyperostotic. For Lake (1898) hyperostosis is distinct from exostosis, being “a more or less uniform bony stricture of the external meatus.” Sabroux (1901, p. 13) distinguishes between exostoses and osteomata, believing the two to differ anatomo-pathologically ; but gives no satis- factory rule for differentiating the two. Korner (1904, pp. 106-7) thus imperfectly distinguishes the two forms: Of the exostoses, the button- to semiglobular forms are found near the antero-superior border of the drum, in the vicinity of the location, still open at birth, of the original tympanic ring. They remain mostly very small. .... The hyperostoses are found as a rule in the outer part of the bony meatus. In accord with the differing participation of the tympanic bone in the formation of the lower portions of the bony meatus, they reach up to the middle, seldom to the upper third of the lumen of the canal. They develop usually synchronously on the anterior and the posterior wall, seldom also on the floor of the meatus, in such a manner that the remaining opening of the canal becomes pear- Shaped)... After what was said it seems that the frequently mentioned occur- rence of multiple exostoses in one ear is really to be assumed as a combination of ex- and hyperostoses; only the small exostoses at the upper border of the drum do occasionally occur in a multiple number oO oreo ic Bezold and Siebenmann (1908, p. 102) offer a somewhat peculiar concept of the hypero8toses and exostoses and decide against speaking of them separately: We call exostoses small round bony growths which are usually found as little white circumscribed elevations in groups of two or more on both sutures of the deepest part of the os tympanicum to the horizontal part of the scale of the temporal bone. One protuberance is usually directly in front, another close behind the short process of the hammer in the drum membrane. A third one often protrudes between the two. Hyperostoses present themselves more in the form of diffuse bulgings of the anterior lower, and sometimes also posterior lower wall. A crosscut through the meatus has consequently the shape of a pear with the point downward. We shall speak about exostoses and hyperostoses together, as they are found not infrequently in the same ear. Jackson (1909) divides the neoplasms merely into: 1, those that entirely block the meatus; 2, those that do so partially ; and 3, those that scarcely affect it, causing merely some narrowing. 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Ballenger (1914, p. 661) attempts, though hardly successfully, to give precise definitions: “An exostosis is a bony tumor growing from the wall of the meatus, and may be either sessile or pedunculated. Hvyperostosis is a diffuse thickening of the bony tissue, or a true hyperplasia.” Ferreri (1904) opposes the division into hyperostoses and exostoses as artificial. But Gray (1910, p. 137) accepts Cassell’s classification, defining ear exostoses as growths “ which are circumscribed and even occasionally pedunculated ; and hyperostoses, in which a general diffuse thickening of the bony walls takes place.” Manasse (cited by Moller-Holst, 1917, p. 71) says that the hyper- ostoses are broadly based thickenings, the exostoses showing a globular development; and the same definitions are given by Bauer and Stein (1926). For McKenzie (1920, p. 457) “ osteomata in the meatus may be sessile (hyperostosis) or pedunculated (exostosis) .” Bauer, Stein, Kaufmann (cited by Alexander) designate the cir- cumscribed protruding bony growths as exostoses, the more diffuse ones as hyperostoses. ‘“‘ Thereby”, says Alexander, “the principal distinction between exostosis and hyperostosis falls; the difference lies simply in the development. The separation of exostoses and hyper- ostoses of the external auditory canal is not easily possible when they occur at the same time in the same ear passage.” Alexander then classifies the growths according to location, into: 1. Exostoses that proceed from the outer border of the os tympanicum. 2. Exostoses that proceed from the inner border of the os tympanicum. 3. Exostoses that arise from the body of the os tympanicum. 4. The flat but slightly protruding exostoses of the inner part of the canal, coming mostly from the terminal line of the os tympanicum. He also classifies the growths pathognomically, of which more later. Oetteking (1930) separates hyperostoses and exostoses, but says that “there may be transitional forms ”, though in shape rather than structure. Finally Moller-Holst (1932, p. 73, 101, 102) expresses himself thus: Under hyperostoses Kessel and Ostmann understand general thickenings of the tympanic bone; Kaufmann diffuse bone increase in mass, but also circum- scribed growths; Hartmann and Virchow flat exostoses; Manasse broadly based thickenings; Marx diffuse malformations which are to be counted among the hyperplasias and which originate already in childhood; Alexander a struc- tural variation, not an abnormality (Missbildung), which when more strongly developed are to be counted with the tumors. Zuckerkandl’s exostoses also, are designated by Alexander as hyperostoses. Under exostoses are generally under- stood localized bony outgrowths. NO. 6 EAR EXOSTOSES—HRDLICKA 45 For Kaufmann, Moller-Holst says, exostoses and hyperostoses are but form-differences. “ Marx, too, knows no boundaries between cir- cumscribed exostoses and diffuse hyperostoses, and no more does Alexander, who uses the two terms for the same growths.” Moller- Holst’s own view is that, though there exist all kinds of transitions between the bony growths in the ears in their form as well as their structure, yet from the clinical point of view there is a justification for subdivisions of the formations on the basis of location, shape, and size. “ Speaking generally, therefore, a so-called bony polyp of the meatus may anatomo-pathologically be an exostosis or a hyper- ostosis ’—which last statement is not very clear. New observations.—It is plain from the above that a generally valid classification of the abnormal bony ear growths has not been realized and in fact, appears to be unrealizable; but that clinically there is a prevalent sense of two or rather three forms—the diffuse or broad- based irregular tume factions ; the circumscribed, occasionally peduncu- lated tumors, capable of a rapid growth and, together with their soft coverings, of causing an occlusion of the whole canal; and small indolent rarely troublesome “ ceiling ” growths in the meatus. The extensive examinations reported in this monograph lead to the conclusion that, aside from the small, more or less pearllike exostoses— or probably better osteomata—arising from the superior or squamous part of the meatus, there is no line of demarcation that would permit any valid subdivision of the growths into hyperostoses and exostoses, or in any other manner. There are many cases where a clear charac- terization of the growth by one or the other of the above main terms, hyperostosis and exostosis, is impracticable—it is more or less both. There are other more differentiated cases in which at the base the growth would deserve the name of diffuse hyperostosis, but above this is definitely a more or less rounded exostosis. There are cases where what is evidently the same growth, in the same location, will in one of the ears of the same skull show the form of a pronounced exostosis, in the other that of some grade of hyperostosis or an intermediary condition. And there are no few instances in which both or all three of these forms (hyperostoses, exostosis, and intermediary) may be seen in the same ear. The more or less diffuse hyperostosis of the pos- terior or anterior wall of the meatus is in many cases in all probability but an earlier stage of an exostosis. Thus otology, for its practical purposes, will doubtless keep its conventional classification of the bony growths of the external auditory canal into hyperostoses, exostoses, and osteomata of the roof; but those who use this classification should be aware of the fact that, 4 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 except perhaps for the rare osteomata of the roof, it is merely a con- ventional and convenient subdivision, without substantial anatomical or pathological foundation. STRUCTURE A number of authors who dealt with ear exostoses report the re- sults of a histological examination of these growths. The results show differences ; some are more or less cancellous and some are compact tumors, but between the two there is no line of demarcation—they connect by intermediary grades. Thus histology, too, fails to afford any definite basis for the differentiation of these neoplasms into dis- tinct categories. About all that can legitimately be said is that the small, pearllike and the smooth, rounded exostoses are generally the more compact, the tumefactions and the large masses prevalently the more cancellous. The structure of ear exostoses received attention from the very first report of these cases, that of Autenrieth (1809). The growth was “ formed of areolar bone tissue.” Contrary to this, in Toynbee’s experience the tumors were compact. In his Textbook (1860) we read: As far as my opportunities have permitted examination, these tumours... appear to consist of extremely hard and dense bone. In one case, where a por- tion of the bone was denuded of membrane, it appeared shiny, white and polished, like ivory. In another, where, under the misapprehension of the body being a polyp, caustic had been applied, the bone was exposed and found to be extremely hard and devoid of sensibility. In a third instance, where I observed the mem- brane to be absent, there was a thin layer of cartilage on the surface, beneath which the bone was very hard. Delstanche (1878) distinguishes “cellular” and “ eburnated ”’ structure of ear exostoses. He says (p. 8): ‘‘ Thus some—the cellu- lar exostoses—are formed by areolar bony tissue which is enveloped by a thin layer of compact bone; others—the eburnated exostoses— are completely formed by compact tissue of much density.” Schwartze (q. by Delstanche) “is of the opinion that the spongy and compact states of the exostoses represent merely different phases of the devel- opment of the neoplasms, and that exostoses that were at first can- cellous can pass into the state of eburnation; and he believes also in the possibility of their transformation in the reverse sense. This opinion is also partaken of by Virchow and Nélaton.” Individual cases were studied histologically by Hedinger, Cocks, Kuhn, Eulenstein, and others. The reports are sometimes rather in- volved, but all agree on one hand in the complete absence of anything No. 6 EAR EXOSTOSES—HRDLICKA 47 that would suggest malignity, and on the other in the lack of a substantial basis for any definite subdivision of the growths into dif- ferent classes. A case of a pedunculated exostosis operated upon and studied by Politzer (1902, p. 210), “‘ showed here and there very compact lam- ellae (eburnation) with spaces that contained but few vessels.” Another pedunculated exostosis was removed and studied by Tod (1909, p. 77) who reports briefly as follows: “ On making a section the ossification was seen to be proceeding from the centre.’ Other exostoses “ were very hard, with a very thin layer of skin over them.” A number of detailed histological examinations of ear exostoses are reported by Dahlstrom (1923, pp. 215-216), and notes on the subject may be found in Kessel, Schlomka, Ferreri, Gray, Oetteking, and other authors, but there is nothing to afford any definite aid in the attempt at a classification of these growths. Marx (1926, p. 502) thus summarizes the results of the histological studies of ear exostoses up to his time: They consist of compact bone without distinct medullary spaces, but often with strikingly marked vascular canals. In substance the “‘ exostosis eburnea ” and the ‘exostosis spongiosa” are not different, but it is to be accepted that one form passes into the other (Weber, Virchow, Schwartze), and that occa- sionally also mixed forms are encountered. Finally, Moller-Holst (1932, pp. 69, 100) states: Some exostoses consist only of cancellous bone and will therefore be desig- nated as exostosis spongiosa, others are covered with a compact layer and are called exostosis eburnea. Histological examinations by the clinicians have shown that between the two extremes (the compact and the spongy exostoses) occur all transitions. Remarks.—tThe only conclusion that can be drawn from the his- tological evidence regarding the bony growths in the ears is that they all together constitute but one abnormal complex, which offers nu- merous individual differences, grades, and variations. NATURE What, in the opinion of those who have dealt with the condition, is the nature, pathologically, of ear exostoses? There are numerous opinions on this subject, but in general they are based on limited in- dividual observations, and that mostly clinical, which do not permit of a sufficient perspective and grasp. However, it is interesting to survey these opinions. Roosa (1866) concluded that ear exostoses “ were morbid growths consequent on local irritation.” 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 For Von Troeltsch (1881) the “ exostoses ” were pathognomically close to tumors, the ‘‘ hyperostoses’’ close to diffuse osteophytic growths. Hedinger (1881) diagnosed his case as “ ostoid metamorphosis of inflammatory proliferation of papillary and connective tissue, with lime deposits *—no regular bone formation. Moos (1881) regarded his cases as simple hyperplasias arising from irritative processes at a time when the tympanic ring fuses with the adjacent parts of the temporal; Roosa, about the same time, viewed them as “ hyperplasias of the periosteum and the underlying bone”; Politzer (1893) as “ partial hyperplasias of development and ossification.” Steinbriigge (1891) terms the outgrowths “ periosteal osteomata.”’ Virchow (1893) considers them as “ plainly products of pathological nature... . examples of disturbed development, which probably begins in and proceeds from the end parts of the annulus tympanicus ”’, and says later, “ The localization of the exostoses of the auditory canal implies that they, as the exostoses of other parts of the skeleton, may be conceived as excessive bone growths on places which are advancing toward ossification.” Whitney puts them, ab origine, under “ cartilaginous exostoses.” Braunberger (1896, p. 6) thought that “if we want to give these exostoses an anatomically and physiologically substantiated position in ear pathology, we must range them with connective tissue tumors in the ectodermal part of the sound-conducting apparatus of the ear.” Ranke (1900) believed that the growths “were examples of an endemic disease common to the burial grounds of Ancon as well as those of old Peru in general.” Sabroux (1901, p. 13) says, “ Exostoses of the auditory canal are tumors formed by an abnormal and circumscribed production of bony tissue on the surface of the canal.’’ They are “ benign tumors, harm- ful only by mechanical obstruction.” Mayer (1923, 1924) attaches these exostoses (as also otosclerosis) to the ‘ hamartomata ”’ or defect-developments (Fehlbildungen), and refers them to a defective condition of the periosteal connective tissue. For Marx (1926, p. 500), “ exostoses of the auditory canal repre- sent circumscribed bony neoplasms and are to be counted with the tumors, in distinction from the more diffuse hyperostoses, that belong to the hyperplasias. A sharp division of the two is, however, not always possible, as evidently there are transitional forms and as cir- cumscribed exostoses may also occur in a meatus narrowed through ‘ ‘ hyperostosis.” No. 6 EAR EXOSTOSES—HRDLICKA 49 Alexander (1930) regards both hyperostoses and exostoses—sep- arable only by their form—when slight as variations, when large as tumors. Remarks—It must be clear from the above that the pathognomy of ear exostoses is a subject that is as yet by no means as elucidated as might be desired. The relative rarity of the condition in the living, its clinical disadvantages, the want in many cases of the assistance of an expert histo-pathologist, all constitute serious difficulties in this field. Perhaps the extensive observations that form the basis of this report may help to throw some light on the matter; this will be dealt with in the terminal discussion. ETIOLOGY The causation of ear exostoses, being of especial importance to otology, has received considerable attention; nevertheless, here also much still remains obscure. The subject has naturally a direct con- nection with that of pathognomy of the growths. A general review of the observations and opinions recorded in this part of the field shows that attention has been concentrated mainly on the local or exciting agencies and on the possible connection of the exostoses with certain systemic conditions or diseases. Never- theless, there are also a number of valuable observations on the heredity of the growths. It will be well to take up separately the various etio- logical factors involved. c TRUE CAUSES “ UNKNOWN ” A large majority of the authors who deal more adequately with the etiology of ear exostoses acknowledge more or less openly that the true or basic causes of this class of disorders are unknown or uncertain. Statements such as “ nothing positive ”’, “ still dark as to the causes ”’, “true causes unknown ”, and others of similar import may be found in Turner, Blake, Garrigou-Désarenes, Ferreri, Lake, and many others. Hedinger (1881), “ pathogenesis entirely unknown”; Whit- ney (1886, p. 41), “ various explanations have been offered for the occurrence of the growth in this situation, but none of these is entirely satisfactory ”’; Kessel (1889), “ causes still but little known ”’ ; Gold- stein (1898), “ etiology often shrouded in mystery ”; Politzer (1902, p. 210), “ the causes of these exostoses are, in the majority of cases, not ascertainable.’ Le Double and Lebourg (1903), “ causation uncer- tain”; Bachauer (1909), “cause not yet found”; Burton (1927), 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 “causes still largely speculative’; Moller-Holst (1932), “ subject in many points not satisfactorily cleared.” Notwithstanding this general pessimistic attitude, there are many beliefs and theories as to the causation of the tumors. They will be reviewed in the following sections. ‘““ SPONTANEOUS ”’, NUTRITIVE There are a number of remarks in otological writings to the effect that an exostosis in the meatus commenced spontaneously without an apparent reason. As late as 1930 Alexander says that some “ originate spontaneously, as an abnormality.” Dupuytren (q. by Sabroux, 1901) suggested a nutritive disturbance. He believed that they depended on a modification in the nutrition of the bone, on some sort of aberration in the distribution of the bone juices, comparing the tumors to the outgrowths which develop as a result of irregularity in distribution of sap. Masini (1882) believed also in disturbed nutrition. He thought the x, 99 growths “ may be comprehended as exaggerated nutritive ‘ lavosio’. CONGENITAL A number of authors have advanced the idea that some of the ear exostoses may be “ congenital’, though none have taken the trouble to state precisely what they meant by that term. In some cases it probably stands loosely for “ hereditary ”’. Thus Dalby (1876) says, “that small bony enlargements in the external auditory canal are sometimes congenital I feel tolerably confident.” Delstanche (1878), a little more explicit, believes that the exostoses “‘ may develop both before and after the ossification of the canal.” For Roosa (1892) “there may be congenital cases.” Velpeau (q. by Le Double) believed some to be “ congenital.” For Goldstein (1898) exostoses in American Indians belong apparently to “congenital’’ growths. Sabroux (1901, p. 24) accepts, though rather loosely, the “ congenital” nature of some of these growths: “we call therefore congenital all exostoses whose origin is unknown and which seem to be transmitted by heredity ; exostoses whose pres- ence coincides with the development of some congenital tumors or malformations; and finally all those exostoses that characterize one race more than another.” In Bezold and Siebenmann’s textbook (1908, p. 102) we read on this subject as follows: “ The exostoses in the bony meatus which Virchow found in many Mexican [should be Peruvian] skulls and which he No. 6 EAR EXOSTOSES—HRDLICKA 51 explained as a peculiarity of the race, must be considered as congenital deformities.” Furst (q. by Moller-Holst, 1932, p. 73) regarded the exostoses as possibly congenital, and Green (ibid.) held that some of the growths could be of fetal origin. All ideas of congenitality are, on the other hand, opposed strongly by Kessel (1889, p. 286), who states: “ In the literature of the subject there are also met with opinions that the exostoses in the auditory canal may occur congenitally. In my estimate this assumption rests on a gross error, for from developmental studies it is known that in the newborn there is as yet no bony meatus and that the same comes into existence only during the first years of life... .. The exostoses are therefore later manifestations. ... . ” Le Double and Lebourg (1903) express the view that the growths “evidently are not con- genital’; and Korner (1904) states unequivocally that they do not occur congenitally. New observations —An examination of numerous skulls of fetuses, newborn, and infants has shown me not the slightest trace of what could be regarded as a hyperostosis or exostosis of the tympanic ring, or on the roof of what was becoming the external meatus. As Kessel pointed out, there is no bony canal until well after birth, in early childhood. Judging from the incomplete developmental stage of the parts up to and for a period after birth, and from the absolutely negative results of our examinations, which included infant skulls of groups that were greatly affected by ear exostoses in later life, it may safely be said, I think, that the growths in question do not occur congenitally. The term ‘“ congenital” should henceforth be excluded from the literature of ear exostoses. DEVELOPMENTAL Aside from causes inherent in the organism and derived either from some constitutional fault or from heredity, a number of students of ear exostoses connect them with “ developmental ” conditions of the external auditory canal. Just what is meant by this is mostly not clear, and even where an attempt is made at explanation, no proof of the cor- rectness of the view is given in any instance. Virchow (q. by Moller-Holst, 1932, p. 73) believed that some of the exostoses originated in the fetal stage from the tympanic ring as a result of a pathological irritation. Much the same view was held by Moos. Green, too, believed that some of the growths could develop in fetal life. For Heiman (1890) congenital ear exostoses were “ due 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 to hyperplasia during the period of development and ossification of the external auditory canal.” Hartmann (1893; 1896, p. 43) held that ear exostoses “ must be cases of anomalies of development, because of the simultaneous ap- pearance on both sides, the limitation of the anomaly to pars tympanica, and the perfectly uniform consistency of the rest of the bone.’ And further (pp. 45-46) : All of these structures are to be regarded as anomalies of formation, which begin with the development of the external meatus and come to light with the advancing of the pars tympanica.... . If I were to offer proof that these alterations were simply anomalies of development and had nothing to do with inflammatory or morbid processes, I should emphasize: the absence of alterations which could possibly be referred to previous inflammation; the perfectly uniform condition of the bone, which can in no way be distinguished from the neighboring bone, and especially from that of the squamous portion; the limitation of the hyperostosis to the pars tym- panica; the uniform occurrence on both sides; the stationary condition of the tumor in adults; and finally the hereditary influence so often observed. Lake (1898) opposes Hartmann’s opinions. But J. Gruber (1897, p. 212) believes similarly “that the development of ear exostoses stands in connection with the growth of the tympanic ring and is to be conceived as an excess product of this.” And for Politzer (1902, p. 210), in some cases the origin of the growths may be attributed to “ partial hyperplasia during the stages of development and ossifica- tion of the bony auditory meatus.”’ According to his view, “ those bilateral osseous neoplasms which develop without symptoms belong to this class, especially if they are located symmetrically in both meatuses and have a corresponding form. They are usually found in the middle and inner sections of the osseous meatus. They are sessile or pedunculated, and seldom attain such a size that they com- pletely fill the auditory canal.” Ballenger (1914, p. 161), too, subscribes to these opinions. He says: ‘‘ They may be due to developmental causes, particularly in those cases wherein the middle and the inner section of the osseous meatus on each side is the seat of the growth. When due to faulty development, the growths are usually small.” Since the above, the “ developmental” origin of ear exostoses received noteworthy further support by Burton (1927). To this observer, it seems logical to believe that the exciting agent, whatever its form, acts equally upon the entire canal but is responded to only by those points which are especially susceptible to the stimulus. Which are these points? In the vast No. 6 EAR EXOSTOSES—HRDLICKA 53 majority of cases reported only those areas responded to this stimulation, what- ever it was, which during the development grew most rapidly. These places are the two ends of the annulus. By far the larger number of aural exostoses are found along the suture lines between the canal and the rest of the temporal bone. Why? The author believes that an adequate explanation is to be found in his theory that these susceptible areas have retained their early potentiality of more rapid development and response to any stimulus of production. For Alexander (1930) the “ deeper causes of ear exostoses lie in the temporal bone, which is formed of parts that ossify in cartilage and parts formed from periosteal ossification; also the persistence of cartilaginous points in the petrosum, and the lateness of the close of the postembryonal growth, create a disposition to variation in the bone and pathological growths.” Critical remarks—Although the theory of a “ developmental ” origin of some, at least, of the ear exostoses has received a considerable backing, yet most of this appears of the order of “ follow the leader ”’. And neither the leader nor those who followed have furnished the slightest material evidence that would bear out their contention, or even make it possible. The discussion under ‘“‘ Congenital Origin ”’ applies also here. There are no observations on record of a fetal, or even later, involvement by the growths up to Field’s child of 3 years, and even that case is unique. The bony parts that are to form the external meatus and from which eventually exostoses may arise, are still largely deficient in fetal life and are still far from complete at birth. And irritative ear troubles, that could excite the formation of the outgrowths, are not known of before birth. INBORN TENDENCY Wholly different from congenitality is the case of the causes com- prised under the term “ inborn tendency ”’. Here is something worthy of earnest attention. The suggestion that there may be behind ear exostoses an inborn tendency toward their production has been expressed by a number of writers on the subject, though one would vainly search for any com- prehensive exposition of the idea. Thus Nélaton speaks of a “ prédis- position générale de l’économie ’’, and Delstanche (1878), who quotes him, admits this as among the causes. Masini (1882) names “ immi- nent predisposition’, and Ostmann (1894) mentions “ inborn ten- dency” as among the causes of the exostoses. For Russell (1900) there 1s a “ tendency in all races towards the formation of bony tumors 54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 or exostoses in the external auditory meatus.” A noteworthy state- ment on the subject is that of Gray (1910): The other general aspect of the problem is that the majority of individuals never do suffer from exostoses, no matter how badly or from whatever cause their general health become affected, or whether they suffer from some local disease of the ear, such as suppuration, middle ear catarrh, etc., or not. Now it appears to the writer that, viewed from this point of view, the logical con- clusion to be drawn is that the cause of exostoses, that is the condition without which the disease cannot occur, is to be found in the organ of hearing itself, and, further, that this condition exists in the organ of hearing of certain indi- viduals, and in these individuals only. From this point of view, which I believe to be the correct one, exostosis is idiopathic. That is to say, the disease occurs in people who have an inborn tendency to it, and that this is the only constant factor in the etiology. No doubt disturbances, either locally in the ears or constitutionally in the bodies of these individuals, may precipitate the onset of the disease, but no particular one of these disturbances need be present and in many cases they may all be absent. The single constant factor is the innate tendency to the disease and the fact that hereditary influences are evident in many cases is strongly in support of this view. Burton (1927) supports Gray’s (and others’) view of the presence of an “innate tendency ” toward these developments, and would call this “atavism”’, though evidently with a different shade of meaning from that ordinarily applied to this term. For Moller-Holst (1932, p. 102) a causative connection of atavism is wholly unfounded. A few authors approach the conception of a “ racial influence ”. Thus Darnach (q. by Alexander, 1930) regards the inclination of the American Indian to ear exostoses as a racial peculiarity (char- acter), something like that of the African Negro toward keloid forma- tions. Moller-Holst (1932) is skeptical and would at most admit a “ race-peculiarity ’ (Rasseneigenthumlichkeit ). Remarks—The above and some additional statements of this nature range from hesitating suggestions to firm convictions. They show that the more obvious agencies of the disorder did not fully satisfy these observers. They felt more or less vaguely, or have become defi- nitely convinced, that some innate predisposition, individual or racial, lay in the background of the etiology of the growths under considera- tion. But not one of them has attempted to pin down and critically analyze the hazy factor. HEREDITY The subjects of ‘‘ congenitality ’, “ predisposition’, and “ inborn tendency ”’, considered above, lead inevitably to that of heredity of ear exostoses, and here otology furnishes a line of definite valuable observations. nN NO. 6 EAR EXOSTOSES—HRDLICKA 55 As early as 1878, Schwartze reports that he had observed hereditary connections, and Delstanche regards “ hereditary predisposition ” as among ‘the best determined causes” of the bony growths in the meatus. According to Blake (1880, p. 88) : there is one other possible influence which may have a bearing upon the occurrence of these growths, and which is at least worthy of consideration in any future investigations which may be made, and that is, hereditary tendency. Of the more marked cases—that is, cases exhibiting excessive growth without evidence of other aural lesion—I have found, in aural practice, that the majority have occurred in certain families, in the male members of successive generations, the most marked instance being in the three successive generations of one family. Masini (1882) found positive evidence of heredity of ear exostoses in one case, and similarly did Boyer, Kirchner, Politzer, and Moore. Kessel (1889, p. 288) states that he can confirm the heredity of the growths out of his own clinical experience. There exists here, he says “‘a constitution-anomaly which leads to certain disturbances in the genesis of the skeleton, without the possibility of a conclusion as yet as to just what this anomaly consists of.” Braunberger (1896) affirms that in individual cases heredity is as- certainable. Hartmann (1897, p. 43) has not only seen evidences of heredity of ear exostoses in his own practice, but a review of the litera- ture on the subject has shown him “the frequent occurrence of hereditary influence in similar cases”; Korner (1904, p. 102) says, “ There exists nevertheless in not a few families an hereditary dis- position to these bony new growths ”’; and in Bezold and Siebenmann (1908, p. 102) we read, “ This anomaly is often found in several members of the same family.” Gray (1910) states that: ‘“ The single constant factor is the innate tendency to the disease and the fact that hereditary influences are evi- dent in many cases is strongly in support of this view.” For Ballenger (1914, p. 662), “ there are some cases in which heredity seems to be a factor in the production of these growths, as the same condition may be traced through a few generations.” O. G. Kessel (1924) declares heredity alone to be the effective cause of the growths, other factors being of but secondary importance. Burton (1927), as already noted, calls the growths “ atavisms”’, which implies inheritance, and asserts that heredity is among the best substantiated of the causes of the abnormalities. For Alexander (1930), ear exostoses are “ often hereditary ”’. ‘ 56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 . Convincing as the above statements are, they are supplemented by the sustaining evidence of actual observations on the heredity of the abnormalities. The reported cases, as far as it was possible to gather them, are as follows: Reports of Cases of Evident Inheritance of Ear Exostoses Blake (1880, 1888): In two families father and son successively affected. Masini (1882): In cne case, adult male, mother “deaf” through occlusion of both ears when about 4o. Boyer (q. by Hartmann, 1897, p. 43): Treated a woman whose father, brother, three sisters, two nephews, and all of her own children were affected by ear “ hyperostoses.” Kirchner (1883): Observed a man with bilateral ear exostoses, in whose son he later found a similar condition. Kessel (1889, p. 288): Like Blake, saw in two families bony growths in ears of a father and a son. Moore (1900): Had two cases, in two brothers, in one in the right, in the other in the left ear. Korner (1904, p. 102): Treated, I, a father, 63, with bilateral marked hyper- ostoses; in his first son, 26, the same; in the second son, 20, the same; in his daughter, 22, a hyperostosis in the right ear; II, father, 55, bilateral marked hyperostosis; son, 7, hyperostosis in the left ear; 1II, man, 42, bilateral marked hyperostoses .... brother 38, the same; sister, 30, meatus normal; their cousin, 19, in right ear a completely occluding, coftee-bean-size exostosis. Kessel (1924): Presents a heredity-tree of three generations of a family with ear exostoses. Alexander (1930): Had three male patients with ear exostoses, the father of each of whom was affected by similar growth; another male whose brother and sister were effected ; and still another male with “ ear troubles ” in a brother and three children of another brother. Discussion.—In view of the above data, it cannot be doubted that direct heredity plays a part in the etiology of ear exostoses. But there is no light as yet on the problems of how large and important this part is, whether it is generalized or restricted to individual families, whether it is progressive or regressive, and how it originates. Also, nearly all that is said relates to direct inheritance in families and not to heredity in a wider sense. The subject will receive due attention in our final chapter. An item that calls for some consideration in this place is that of inbreeding. Inbreeding as a cause of ear exostoses was brought up recently (1932) by Moller-Holst, but without comment or any attempt at an explanation. Inbreeding is known to be capable of accentuating and spreading different pathological conditions. In limited localities, in small human groups, and especially in single families, it might have No. 6 EAR EXOSTOSES—HRDLICKA 57 played a part in multiplying the cases of ear exostoses. But inbreed- ing means essentially intensified heredity. It could augment but hardly create the exostoses. As these abnormalities are so widespread over the earth, inbreeding, if it was their cause, would have to be equally generalized, which of course it is not. Harmful inbreeding is rare in any race of men. It did not exist in America, where exogamy was the rule. It did exist in Egypt, but only in the Pharaohs and perhaps in the highest families. It took place only very exceptionally in whites. More closely inbred communities, whether in the Old World or the New, have never been reported as especially subject to ear exostoses. In fact, the more closely the subject of inbreeding is examined the more it eludes as a possible cause of ear exostoses. SOCIAL STATUS: OCCUPATION There are but a few references to social or occupational causes of ear exostoses, but these few are suggestive. Von Troeltsch (1881, p- 142) states that his cases predominated in those of middle years ‘“ who liked to eat well and drink.” Bezold in 24 years of hospital and private practice found ear exostoses in 116 patients and says (1895, p. 49): “In only a few instances have I found the exostoses in the dispensary patients; my observations come almost all from private practice. The occurrence of the exostoses is therefore over- whelmingly a disease of the better situated.” And the same is reas- serted in Bezold and Siebenmann (1908, p. 102). Korner (1904), in dispensary practice at Rostock, found ear exos- toses in but 1.1 percent of his ear patients—a relative rarity in the poor. For Tod (1909) the growths appeared more frequently in the “more favorably circumstanced than in the labouring class.” Moodie (1930) mentions that the exostoses are more frequent “ among the sedentary groups than among the more active ’’, although he does not state the basis for this conclusion. In Moller-Holst (1932, p. 69) we find the following statement, though it is perhaps not original : “Usually [the ear exostoses] will, curiously, be found almost ex- clusively among the well-to-do and but seldom in the poor.” Remarks.—Both the social and the occupational factors are of such breadth and complexity, as well as potency, that some influence by them on any pathological or abnormal process may safely be assumed. Such influence may be of direct or indirect nature, primary or secon- dary. Given the existence of a widespread oto-exostotic diathesis, it is conceivable that its manifestations might be favored or hindered by what is embraced in the term “ social status’, or occupation. Either 58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 of these complex agencies, in other words, might act as a contributory, exciting, modifying, or checking cause of the abnormalities under consideration. CLIMATE The climate of England has been blamed. Law (1909, p. 77) thought ear exostoses were ‘“ more frequent in the South, and more so among individuals who have lived in hot climates.” Alexander (1930) regards climate as possibly influential in “ releasing” the growths. Moller-Holst (1932, p. 69) says—though apparently not as a result of personal observation—that “it is also an interesting fact that the development of exostoses in the external auditory canal occurs more frequently in people who had had a prolonged stay in the tropics ’’; yet on another page (102) he states that this view is “unfounded ”. Remarks —Our materials hardly sustain the claim for climate as one of the causes of ear exostoses. Our northernmost large group, the Eskimo, is nearly free from the growths; but so is one of the most southern groups, the Egyptians, and so are even more, apparently, the essentially hot-climate African Negroes. The exostoses are fre- quent along the coasts in Peru, but so they are in the highlands of Peru and Bolivia. The old tribes of the Channel Islands in California and those of Florida, who were equally at least as “ oceanic ” or litoral as the Peruvians, show much less incidence of these formations. This is particularly noteworthy in Florida, whose old natives belonged largely to the same physical type as those of Louisiana and Arkansas, and where moreover there was a prevalent tendency toward a super- abundance of bone formation, as shown frequently by all parts of the skull as well as the skeleton. The greatest involvement by the ex- ostoses is found in tribes of Dakota, Kentucky, Virginia, Tennessee, and Arkansas—all inland regions but differing considerably in cli- matic conditions. And there is the considerably greater frequency of the growths in the Polynesians than in the Melanesians, with whom the climate is very similar. From the above it seems that for the present the only safe con- clusion to be drawn from the geographical distribution of tympanic exostoses is that their frequency differs very considerably in differ- ent territories, but that this is largely, if not entirely regardless of climate or other geographic factors. FOOD, DRINK, HABITS Stimulating food and drink.—Toynbee (1860), aside from other considerations, says: ‘‘ The disease in question may be divided into No. 6 EAR EXOSTOSES—-HRDLICKA 59 two classes: I, appears with congestion of the mucous membrane of the ear. Most of the patients who have consulted me on account of it were in the habit of partaking freely of stimulating food and beverages.” Hedinger (1881), on the basis of about 40 cases, found it necessary to deny abuse of liquor as an etiological factor. Heiman (1890) mentions alcohol, but the question is ‘ hard to decide.” Ac- cording to Alexander (1930), food has possibly also “‘a releasing effect on the exostoses.” Moller-Holst (1932, p. 102) says “ effect of nutrition is not determinable.” Habits —Habits in the main imply occupations. With the possible exception of those of sea divers (see under Salt Water), and that of carrying heavy ear pendants (see under Mechanical) no attempts have thus far been made to connect occupational or other habits with ear exostoses. CONSTITUTIONAL DERANGEMENTS GOUT, ARTHRITIC DIATHESIS In dealing with the obscure etiology of ear exostoses it was in- evitable that much thought should be given to the possible connection of these abnormalities with those constitutional derangements or dis- eases that are known to be the cause of osteoblastic disturbances. The foremost of such conditions are the arthritic or gouty diathesis, and syphilis. The view that rheumatic or arthritic diathesis and resulting localized osteitis or periostitis were the causes of ear exostoses was suggested as early as 1856 by Rau, but was especially fathered by Toynbee (1860), who believed that “it 1s principally in individuals of rheu- matic or gouty temperament that one encounters these bony tumors ” ; and “ they appear to be the result of a rheumatic or gouty diathesis.” Roosa (1866, p. 428), is the first to raise a partial objection—some influence of rheumatic dyscrasia cannot be denied, but must not be overestimated. For Delstanche (1878) the influence of rheumatic or gouty diathesis is not well established. Von Troeltsch (1881) did not see arthritic signs in his cases. For Dalby (1885) the views of Toynbee and similar views are “ fallacious’’; for Kessel (1889), ascribing the growths to rheumatic or gouty diathesis is untenable. Virchow, (1889, p. 395) is uncertain—“ there exists a certain re- lation between these formations and arthritis deformans’”’; and in 1893 he is still in doubt—there is “ possibly some connection.” Jacque- mart (1889, p. 192), however, returns to the old view and believes ear exostoses result “ from the species of plethora which characterizes arthritism; are found principally in individuals of rheumatic and 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 gouty temperament.” Pritchard (1891), speaking of ear exostoses of certain kinds, says “as far as my own observations go, I am inclined to the belief that they are usually of gouty or rheumatic origin.” Alderton (1898) saw in his case “a local gouty manifestation of the constitutional diathesis.” For Politzer (1902) gout was one of the causes of the tumors; for Tod (1909) one of the chief causes; and Noquet (1899) saw a case of bilateral ear exostoses in a gouty subject. Sabroux (1901, p. 27) sums up the situation as follows: ‘“ Aside from the syphilitic diathesis, we must mention in the etiology of these conditions, arthritism and herpetism, diatheses which are nearly equivalent and closely related if we consider their effects, and finally the gouty diathesis.” For Ballenger (1914, p. 662), gout is doubtful as a direct cause; “it is more probable that the gouty diathesis causes an inflammatory process of the skin and the periosteum, which finally undergoes a retrograde change and becomes the seat of lime deposits.” Remarks.—Arthritic, gouty or rheumatic diathesis is a subject still far from being well understood. It is not even definitely known whether all the conditions comprised under these terms are merely so many differing manifestations of one basic diathesis, as it seems to the writer, or whether they belong to two or more related yet dis- tinct constitutional derangements. They do not, except perhaps on very rare occasions, affect the external auditory canal. They cause irregular osseous deposits in or about the joints, but never peduncu- lated or circumscribed bony tumors. And they affect mainly those above middle age, whereas exostoses are predominantly an affliction of the earlier part of adult life. A very large proportion of the rheumatic, arthritic, or gouty, moreover never develop ear exostoses. Where then is the connection between the two conditions ? That ear exostoses may exist or develop in “ gouty ” subjects need not be questioned ; that a gouty condition of the system may possibly in some cases act favorably on their development could readily be admitted, though proof should be furnished; that an arthritic di- athesis could be the sole cause of an ear exostosis is unexplainable and is opposed by the mass of evidence presented by the extensive skeletal materials at our disposal. The Eskimo offer perhaps the best refutation of a causal connec- tion between arthritis and ear exostoses. Arthritis is frequent and at times severe in these people, leading to marked joint deposits and marginal exostoses—ear hyperostoses and exostoses are practically absent. On the other hand, in our Kentucky, South Dakota, and Vir- ginia groups, where ear exostoses are most common, arthritic condi- tions are no more frequent than usual in American material. No. 6 EAR EXOSTOSES—-HRDLICKA OI SYPHILIS Syphilis has been blamed for so many pathological conditions of which the causes were difficult to detect that no surprise need be felt that ear exostoses, too, have been attributed to the disease. The first to believe he saw a direct connection between the two is Triquet (1857, q. by Sabroux), who says “ the presence of these bony tumors is not encountered except in subjects manifestly affected by syphilitic infection.” Roosa (1866, p. 428) believes that, as with gout, “the influence of syphilis may not be denied, but must not be over- estimated”. Gruber (1870) thinks syphilis acts in part as a cause ; so also, more or less similarly, do Politzer, Jacquemart, Fournier, Noquet, Meniére, Krakauer, and Sabroux. To the above stand opposed Von Troeltsch, Delstanche, Schwartze, Erhardt, and Hedinger. Von Troeltsch (1881, p. 142) says cate- gorically—‘“ connection with syphilis is utterly undemonstrable ”’. Hedinger (1881), as a result of his observations on about 40 cases of ear exostoses, “ must exclude syphilis as an etiological factor”. For Kessel (1889), ascribing them to syphilis is untenable. According to Duplay (q. by Sabroux, 1901, p. 25), “ nothing authorizes us to connect these exostoses with syphilis.” Sabroux (1901, p. 24), though on the basis mainly of the opinions of others, inclines to the view that syphilis is concerned in the genesis of the congenital ear tumors; while in the acquired forms, “ syphilis is quite certainly the most common agency that influences their de- velopment. .... In hereditary syphilis we have the congenital ex- ostoses; in the acquired we encounter the exostoses as one of the tertiary manifestations of the disease.’ And for Ballenger (1914, p. 662), ‘“‘ Syphilis is undoubtedly a cause of the growths, although not in a very large number of cases.” Yet for Bezold and Siebenmann (1908, p. 102) “ there is no proof of a special diathesis for this disease such as lues.” There are other statements on the two sides of the question, but no proofs. More recent textbooks and authors in otology tend to be cautious. Remarks.—Not one single case in the great collection of prehis- toric American material that passed through our hands in connection with this study, has shown any evidence of syphilis. Nor was there seen any trace of the disease in any of the Polynesian or other skulls that were found to be affected with ear exostoses. In the 14 post-Columbian to recent Indian, Eskimo, and White skulls with extensive tertiary syphilitic lesions, in the United States National Museum collections, not one shows even a small ear exostosis. 5 62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Whether syphilis, acquired or inherited, may in some way favor the appearance or growth of ear exostoses cannot be decided from my sources. But these do show beyond question that ear exostoses may exist, and that to a far greater extent than in White people, without a trace of syphilis being present; and that on the other hand a whole series of skulls with most pronounced syphilitic involvement may exist with not a vestige of ear exostoses. LOCAL IRRITATIONS AND INFLAMMATIONS Of all the possible causes of ear exostoses, none in otological litera- ture receives as much attention as ear irritations and inflammations. The irritations are secondary, due particularly to injuries and dis- charges, and act by setting up an inflammation. The inflammations may be of any intensity, acute or chronic, localized or generalized. They range from mild focal periostitis or osteitis to serious involve- ments of the meatus, and to otitis media. All were, and largely still are, believed to be more or less directly influential in favoring the development of the bony growths in the meatus, directly or through their irritating discharges. Wilde (1855, p. 241) concluded that ear exostoses were the results of chronic osteitis and periostitis in the meatus. Rau (1856) ex- pressed the same opinion. For Toynbee (1860), one class of the growths was due to “ congestion of ear lining”, secondary to a dis- eased condition of the ear. Roosa (1866, p. 428) saw in their pro- duction a process “ substantially an irritative one, often even inflam- matory’; “irritation and probably periostitis due to chronic otor- rhoea.”’ Similarly for Dalby (1876) they were “ at times called into existence by an irritation .... by the irritating influence of a discharge ”’. Delstanche (1878, pp. 17, 65) enumerates ‘ among the most potent causes ” of ear exostoses, “‘ chronic phlegmasia (inflammation), pri- mary or secondary, of the walls of the meatus . , . . inflammation of bone and periosteum ... . inflammation of irritation, spontan- eous or traumatic’. Similarly, the chief or one of the chief causes of the growths is, in the opinion of Hedinger (1881), “ hypertrophic inflammation of the lining membrane, with subsequent osseous meta- morphosis of the new-formed connective tissue.” For Masini (1882, p. 616), continued local irritation (as in otorrhoea) is a cause of the growths. Moos, Kessel, Politzer, Pritchard, Jacquemart, Heiman, Kuhn, Jackson, Sabroux, Ballenger, Michailowskij, and others express themselves in a more or less similar manner. No. 6 EAR EXOSTOSES—-HRDLICKA 63 xX ‘ For Roosa (1892) the growths are “in general caused by local irritation.” Politzer (1902) sees as their basis hypertrophic inflam- mation of the lining of the canal, with ossification of the new tissue. Virchow (1893) feels that they “ owe their inception to a pathological irritation restricted to the pars tympanica.” For Green (1896) they are “in most cases, in all probability, the results of a circumscribed periostitis.” For Goldstein (1898) “the most tangible and compre- hensible cause, yet one not often met with, is that of the long-standing direct irritation and chronic inflammatory condition of the walls of the external auditory canal.’”’ McKenzie (1920, p. 458) holds that “exostoses of the meatus seem to be the result of some long-continued irritation, such as chronic suppuration of the middle ear, or eczema of the meatal wall.”” For Michailowskij (1924, 1926) the chief cause of ear exostoses is chronic otitis media and long-lasting dermatitis of the auditory canal. Ballenger (1914, p. 661) tries thus to explain the process: “‘ Chronic suppuration of the middle ear may excite a secondary inflammation of the membranous canal, and cause a fibrous or connective tissue thickening, which, after a long period of time, may be metamorphosed into osseous tissue.” Many other authors, however, express doubt. Field (1878), al- though acknowledging the influence of inflammatory conditions, never- theless points out that there is a “ rare form of exostosis of ivory con- sistency, partaking of- the nature of a new growth, and quite inde- pendent of inflammatory changes.’ Kessel (1889, p. 287) and after him Schlomka (1891, p. 15) acknowledge that the exostoses occur most frequently in those with chronic ear discharges, and that the discharge causes irritation is beyond doubt; but often it is very hard to decide which was the first condition, for it can be shown that the exostoses are capable of arousing inflammation. Bezold (1895, p. 50) found that ‘‘ suppurative processes of the middle ear, as accompani- ments of exostoses, belong to the great rarities.” Korner (1904, pp. 102-104) feels that the causation of the hyper- ostoses and exostoses of the meatus by other ear affections is a pos- sibility only in very isolated instances and definitely proved in none ; but ‘‘ there is a rare variety of the exostoses that appears only in con- sequence of chronic suppuration. These are small buttonlike, fairly pedunculated growths that develop from the postero-superior border of the meatus, close in front of or behind the suprameatal spine and which therefore do not belong, as do the ordinary exostoses, to the tympanic part, but to the squamous portion of the temporal bone.” Gray (1910, p. 137) cautions that “ judgment must be reserved in this matter’’-—a large number of the growths are without any symptoms. , 64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Burton (1927) is skeptical as to the influence of otic discharges: “If the exciting agent be some form of chronic irritation and the potentialities of all four walls be equal, we are surely entitled to postulate that the floor, being the most dependent part, would be the wall receiving the maximum stimulation, and hence should be the area most frequently productive of these overgrowths. But the floor is hardly ever involved in the process. It follows therefore that we are justified in premising etiological factors other than chronic irri- tations ue, ” O. Mayer (q. by Alexander, 1930) does not believe that ear exostoses arise from a primary local inflammation. Moller- Holst’s material (1932, p. 102) “ fails to show connection with otitis media ”’; and he has also failed to find, in the mummies with ear ex- ostoses, inflammatory processes in the soft parts. Discussion.—As with most other alleged causes of ear exostoses, the question remains undecided. There is no question but that in a large majority of the clinical cases the growths were found accompanied by otorrhoea and in some by marked otitis media. But there were also observed cases without any discharge or otitis, and a great many patients with otorrhoea or otitis media develop no ear exostoses. A number of the aural surgeons reporting on the condition mention that for long periods the growths caused no trouble to the patient. In general the subject is unaware of the hyperostosis or exostosis until it reaches such a size that it begins to be accompanied by a discharge or interferes with the hearing. But when conditions reach such a stage, a determination of which was first, the bony growth or the discharge and inflammation, is impossible. The fact that the growths almost never develop from the floor of the meatus, which is most exposed to ear discharges, need not have much significance, for in all probability, owing to differences in in- nervation and blood supply, not all parts of the tympanic bone are equally prone to the development of exostoses, and at night the dis- charges bathe also the lateral parts of the meatus. Theoretically, 1t cannot but be acknowledged that any cause in- ducing nerve irritation and reactive prolonged hyperaemia of the meatus would be capable, where a “ predisposition’ to ear exostoses existed in the parts, of arousing or favoring their development. Where the predisposition does not exist or is held in check in some way, there would, in all probability, be no exostoses. Irritations and inflamma- tions may thus be admitted as among the possible exciting causes of ear exostoses, but hardly as the basic causes of such growths. NO. 6 EAR EXOSTOSES—HRDLICKA 6 Sal The Indian and other material that passed through my hands sus- tains the view that the fundamental causes of ear exostoses are quite different from mere irritation or inflammation. Most of the specimens with such growths in our collections show no detectable inflammatory changes. Nor are the growths themselves, in a large majority of cases, such as would suggest inflammation. They and the parts about them show nothing like the periostitic bone deposits or osteitic indurations that are so well marked elsewhere in the skull or skeleton where inflammatory processes affected the bone. Our material suggests strongly that, in general, otorrhoea and other otic derangements may not precede but rather follow ear hyperostoses and exostoses, after these have begun to reach an obstructive stage. SALT WATER Connected with the preceding category of causes, yet having some individuality of its own, is the irritation of the ear by water and more particularly salt water. A number of authors incline to regard this as a cause of ear exostoses in some instances. Thus in Field’s (1878) cases, two of his four patients attributed their ear troubles, which proved to be exostoses, to repeated and pro- longed sea bathing. For Korner (1894) exposure to sea water ‘‘ was influential ”—he found them more than five times as common along the seacoast of Germany as in the interior. Moore (1900, p. 786) re- ported that ear exostoses were “‘ very common in the Hawaiian Islands, where great fondness for aquatic sports is supposed to be a cause.’’ Tod (1909) blamed “ gout and the morning tub.”’ For Law (1909, p. 77) “a causative factor seemed to be the pouring of water into the ears, especially carelessly sponging while holding the head on one side ’’; and for Jackson (1909) it was “ local irritation, principally by sea water.’’ Jackson thought irritation by salt water to be a more frequent cause of ear exostoses than any other. McKenzie (1920, p. 458) says, “they are very common in people who indulge freely in sea-bathing and diving.” For Heiman (1890) the case is “ hard to decide.” In a case of Lake (1898) the subject “had never had a sea bath.” And for Moller- Holst (1932, p. 102) the claim, with some others, is “so unfounded that no discussion of it is necessary.” Remarks —Some efforts were made to connect the frequency of ear exostoses in the Peruvians and the Channel Islands Californians with sea bathing. It was not known then that the abnormalities would be found to be even much more frequent in some of the inland tribes; that they also occurred in parts of Peru where no one bathed ; that they 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 were rare in the Egyptians who were great water lovers; and that the Malays and the boys of some other maritime groups, including certain Whites, were great swimmers and divers without being par- ticularly troubled with ear exostoses. Notwithstanding the above, sea water is undoubtedly capable of causing chronic ear troubles, and these may conceivably act, where there is a tendency to the exostoses, as exciting ‘or favoring causes. ABSCESSES, FURUNCLES Cassells (1877) regarded some ear exostoses as secondary to a subperiosteal abscess of the mastoid. Jacquemart (1889, p. 193) states that after a cure of an ear furuncle or abscess, it is not surprising to see a development of one of these tumors ; and Sabroux (1901, p. 29) makes the same assertion. POLYPS From evidence adduced by others, Kessel (1889, p. 289) is satisfied that ear polyps may ossify and thus turn into bony tumors. Pritchard (1891) attests that the origin of ear exostoses in some cases may be traced “ even to the actual ossification of the polypi.” Sabroux (1901, p. 31) quotes Klotz, Bezold, Patterson, Cassells, Hedinger, and Cook as having seen bony tissue in ear polyps, seemingly parting from the bony part of the canal and developing preferentially into the im- plantation of the tumor (Politzer). Tod (1909) reports a case where a single exostosis developed “‘ from the floor of the auditory canal ”’ 2 years after the removal of a polyp from that ear. Dahlstrom (1923, p. 216) reports that ossification of ear polyps had been observed by Cocks and Noltenius. Remarks.—These cases, if correctly reported, would seem to differ substantially from ear exostoses. MECHANICAL CAUSES AND TRAUMATISM The first to call attention to the probable influence of mechanical causes on the development of the ear exostoses was Seligmann (1864), who advanced the theory that in ancient Peru the growths were due to the mechanical irritation produced in the Incaic youth, about the time of puberty, by loading the ears with heavy ornaments. This idea is met with also here and there in later authors, and even Virchow (1889, p. 395) inclines to the view that such may have been the cause of the growths in some cases. Blake (1880) found it sugges- tive that most of the growths occurred in the posterior wall of the meatus, “ the wall most exposed to violence.” No. 6 "EAR EXOSTOSES—HRDLICKA 67 Politzer believed traumatism might be a cause of ear exostosis, through consequent localized periostitis. Jacquemart (1889, p. 193) claims that even in cases of arthritic or syphilitic diathesis which pre- disposed to the bony growths in the ears, “ it is ordinarily by trauma- tism that the process commences.” More or less similar views are advanced by Heiman (1891), “L. B. of Hamburg”’, and other authors. Wagenhauser (q. by Schlomka, 1891, p. 14.) reports a case in which, following a fracture of the anterior wall of the meatus resulting from the kick of a horse, 3 months after the healing of the wound there showed on the site of the injury in the canal two small rounded exostoses. Contrary to these opinions and reports Kessel (188 9) declares that “should one think of mechanical irritation, he will be disap- pointed. We see foreign bodies remain in the auditory canal for whole decennia, without causing exostosis.” Sabroux (1901, pp. 24, 28) tends to restore the belief in such causes ; he says: A purely mechanical cause may also determine deformations of the auditory canal and thereby bring about the production of exostoses. Such an agency might be the compression of the external ear by the umbilical cord when this is coiled about the head..... An imjury of the meatus, however slight, may become the starting point of an exostosis, particularly in those predisposed to such growths through syphilitic, rheumatic, or gouty diathesis; and the same applies to severe traumatisms. Korner (1904, p. 104) says that “also fractures of the temporal bone, which pass through the auditory canal, may lead to the de- velopment of hyperostoses or exostoses in the canal. I have seen two such cases. In both the new bone formation assumed the form of a dull cone and was located on the floor of the canal quite far out toward the mouth of the meatus.” Ballenger (1914, p. 661) holds equally that “traumatic fracture of the walls of the meatus, whereby a circumscribed periostitis is excited, may finally result in the formation of a bony mass or tumor.” Moller-Holst (1932, p. 102) objects to the presumed influence of ear pendants. There are still other references to the subject of traumatisms in otological literature, but they add nothing further. From the above it appears certain that traumatisms of the meatus may in some cases act as the exciting causes for the development of bony growths in the canal. On the other hand no substantial support 68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 is given to the idea that heavy ear pendants might induce the de- velopment of these growths, though conceivably in rare cases some exciting influence may exist even in this connection. HEAD DEFORMATION Students of ear exostoses who gave attention more especially to American cranial materials, where artificial deformation is frequent, have mostly been inclined to attribute to these deformations more or less influence in the causation of the abnormalities ; but there were also those who opposed the view. There are three main forms of head deformation. One is the simple occipital flattening produced by the unaided or aided pressure of the back of the head on a resistant base. This form assumed its greatest development and vogue among the Pueblos, but it was also present in Florida and elsewhere. The next form is the “circular” or ““Aymara”’ deformation, pro- duced by a band applied about the head from over the forehead to under the bulge of the occiput. This caused the curious more or less truncated skulls known best from the highlands of Peru and Bolivia, but common also in Vancouver Island, in parts of Argentina, and in other localities. The third form of head deformation is the fronto-occipital or ‘ flat-head” compression. This was produced by the direct application of pressure to the forehead by means of planks or bags, and occasion- ally a pad or other appliance for counterpressure to the lower part of the occiput. It prevailed in the Columbia basin, in the south- eastern United States, in Mexico and Yucatan, in northern South America, and in Peru. It was also present in Hawai. The presumed influence on the development of ear exostoses of artificial skull deformation, ranges itself with the mechanical causes. About the best expression on the subject is that of Whitney (1886, pp. 441-442). He says: If now the flattened skulls are examined it will be found generally that the meatus is narrowed from before backwards and the lips are often slightly thick- ened and raised up; .... There is no absolute demonstration possible that it is the narrowing of the meatus from posterior pressure in early youth that gives a vicious twist to the tympanic ring and places it in a condition favorable to give rise to such outgrowths in after years, All that can be said is that it occurs more frequently in such heads than in those that are normal or flattened by anterior pressure which does not apparently affect the shape of the meatus. And further the similarly flattened heads of the ancient Peruvians show also a large percent affected with hyperostoses. It is not claimed that this deformity is the sole cause, but that it simply increases a tendency which is universal. NO. 6 EAR EXOSTOSES—HRDLICKA 69 For Ostmann (1894), too, the cause of the tympanic exostoses among the American aborigines 1s largely connected with the deforma- tions of the skull, through its effect on the meatus, which sets up ir- ritation that will lead to exostoses, especially when any other tendency toward such formation exists; with Whitney, he recognizes in the American materials two main causes—‘ inborn tendency, with head deformation.” Goldstein (1898) inclines to a similar opinion ; Russell (1900) believes the tendency to bony tumors in the meatus “is in- creased in deformed crania’’; and there are other suggestions of this nature. On the other hand Turner, as early as 1879, though he found exostoses in a deformed Peruvian and a flat-head Chinook, states that “ there was nothing in the two skulls to indicate that the growths may have been induced by the artificial deformation.” For Virchow (1893) head deformation was without effect on the production of these growths; for Ten Kate (1896) head deformation had “ no con- nection with ear tumors ”; and for Von Luschan (1896) ear exostoses “cannot be attributed to head deformation.” It is plain from the above that the influence of artificial skull deformation on the production of ear exostoses is still problematical. However, with the observations already on record and the extensive materials that form the basis of this study, it should be possible to throw more definite light on the matter. Let us see how the principal American data on ear exostoses will range themselves in relation to head deformation. The lessons of the table on page 70 are not as conclusive as might be desired, yet they are not valueless. The records show that undeformed American Indian skulls may be practically free from ear exostoses, may be involved slightly to moderately, as in the Californians, or may present a very marked involvement, as in the Coahuilas. An absence of cranial deformation is therefore no index of freedom from the growths. There appear in the table several other points worthy of notice. The Argentine Calchaqui, though generally affected by more or less of occipital cradle-board flattening—artificial but very gradual and prob- ably unaided deformation—are, in the available series, free from ear exostoses ; and the Vancouver Islanders, all of whose skulls show the artificial circular or Aymara shaping, which in some cases 1s known to reach very marked degrees on the island, show the growths relatively but very rarely. On the other hand the Chinooks, all of whose heads are deformed by more or less pronounced fronto-occipital flattening, present a very high proportion of the abnormalities. 7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Older Data on American Indians Number| _ Per- ae of skulls}centage Author Territory Deformation o cae S!with earjof skulls rae ex- |with ex- amined ostoses | ostoses Oetteking....| NW. Coast, NONE. soe eee eae 112 misc. Ten Kate....| Calchaqui..... generally occipital 110 flattening Russell...... Californias ces NONeGs. meee eee 158 2 ie Oetteking....| Vancouver all deformed, circular. .| 260 5 Ii. Island Russell...... enue ciysns oes most deformed, fronto- | 447 | 24 5. occipital & circular Alexander....| Amer. Indian, in general, no 550 | 30 5. chiefly deformation California Wirchowme a. ANCONA Ee most deformed, fronto- | 134 18 136 occip. Rousselle yy. Tennessee & most deformed, fronto- | 456 69 15. Ohio occip. Moller-Holst | Chile-Bolivia....| most deformed, fronto- | 341 57 1 Ge9/ occip. Oetteking....| Chinook........| all deformed, fronto- S20 2am ela occip. Studley... Coahuilaee NONE Se ca ree 22 a Git ote: New observations —In the next table are given the results of our own series. The indications from these data are hardly better than— in fact, not very different from—those that preceded, but there are confirmations which can hardly be without importance. In the first place it is now definitely seen that undeformed groups may range widely in the incidence of ear exostoses. This can only mean that the syndrome of these growths exists independently of cranial deformation. The second synergistic result is that shown by the old Pueblos. A large majority of the skulls of this large group are deformed by the occipital cradleboara compression. In some cases this compression seems to have been aided, is very pronounced, and the whole skull has been affected as a result. Yet this group shows a remarkably low pro- portion of cases of the abnormal ear growths. It thus ranges itself with NO. 6 EAR EXOSTOSES—-HRDLICKA _ Fis the Calchaqui, who present the same variety of deformation. These results are further strengthened by Hooton, who failed to find any noteworthy involvement in his large Pueblo series from old Pecos. All this justifies the conclusion that the simple occipital cradleboard compression does not favor—and may possibly even check in some way—the development of ear exostoses. On the other hand the fronto-occipital artificial deformation, pro- duced by the direct and forcible application of pressure over the fore- head with a counter pressure on the back of the head, must be viewed with the increasing suspicion that it aids in some way in the ap- pearance of the growths under consideration. Author Hrdliéka New Data Numb f ‘ Aa Be sculls Territory Deformation 2 eels S)with ear amined eataees American: ESS kettria Operas ye eae loiccaetc eNOoee a ST I ,000 2 Old Pueblo........ nearly all deformed, BOG) 212 occipital flattening INEM Akola ae Sacer oie etme Alia aig | 29 2 Mlonidane 5 86 about 10 percent 205 | 35 deformed, fronto- occipital Caiarmice re sei sh tit eae sean es cake 435 | 46 INES States sey). |semeinerise 1 cone ae 112 13 Rens AeA RG cot ors most deformed-mainly |3,651 | 522 fronto-occipital— some circular WANS) (os Rae en ae gel [Oe Nee Ce, ord ee a 65 14 Wouisianas seen - frequent deformation, 61 15 fronto-occipital Arkansas.......°..| occasional deformation,| 173 47 fronto-occipital Ss Dakota alts ot .5| cotnon nas cove comitocecs sete 109 30 Ss DakMowbridgeliias .csce ee ae se oe hes 76 22 Kentuckyaanianiae- only a few deformed, go | 29 occipital or fronto- occipital Polynesian: Weir 2 CALAHOE sree muctoide Qathieice sear aih cs bces 19 4 Hla ware past sree most undeformed— 148 39 occasional fronto- occipital Per- centage of skulls with ex- ostoses No oo AND This matter may perhaps be more conclusively tested in another way. In six of our series exact records were made of the three main kinds 72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 of deformed skulls (fronto-occipital, circular, and simple occipital), and their aural exostoses. The results came out thus: Frequency of Ear Exostoses in Undeformed and Deformed Skulls of the Same People, and in the Three Varieties of Deformed Percent of skulls with ear exostoses in— erritory nee undeformed | inital | Circular or | gether BCT eerie een verona hin eae, eens (O57 14.4% Be 18.0 Pele DlOStee a wae oiovs ees hats 2.0 Dias d. absent | d. absent WoUtsicima area dee eecks! ie cists tas 21.7 detcrmation absent 26.3 NT KANSAS wee oe eee Chios einem see 21.6 absent absent 31.8 Hlonidamew erm ste cec ce 8.7 absent absent 12.5 IGEN Gs 'ordta 0 cyclic Oe HRS eee 23.5 absent absent 28.3 2 Most, if not all, of these Peruvian skulls are really cases of artificial ‘‘flat-head’’ deformation, but the pressure on the forehead was not sufficient to cause a permanent well-marked flattening of the frontal bone. In this table, it will be noted, it is possible to compare directly the percentages of specimens with exostoses in each category of the unde- formed and deformed skulls, within the same ethnic series. The re- sults are convincing. In every instance where both undeformed and “ flat-head ”’ skulls existed in some quantities, the proportion of ear exostoses is distinctly higher in the flat-heads. The fronto-occipital deformation, therefore, it may now for the first time be said posi- tively, favors somewhat the development of the growths. The simple occipital or cradleboard flattening has evidently but small if any influence on the exostoses ; and the circular or Aymara artificial deformation seemingly shows more of a checking than a stimulating effect on the growths. Just how the flat-head deformation may favor or excite the develop- ment of ear exostoses is not clear. Many of these cases of deforma- tion are not of extreme grades, and the skulls that are most deformed do not by any means always show the abnormal ear growths; the occurrence of the latter, in other words, is not directly proportionate to the grade of the distortion. Where the occiput has been much flattened in these cases—pressed forward—a considerable stress has undoubtedly been transmitted to the parts composing the external bony ear. But much the same com- No. 6 EAR EXOSTOSES—HRDLICKA 73 pression of the back of the head with similar effects on the ear occurs in some of the cases of the simple occipital flattening, without resulting in tympanic exostoses. It seems that it is the forcible frontal compression which is mainly to blame. The stresses within the skull produced by the fronto- occipital forcing are certainly greater than those in simple occipital compression. The shape of the external auditory canal is certainly affected in many of these cases. But just how this could give rise to the exostoses is not clear, especially as such growths do not appear to develop earlier in the flat-heads than they do in other crania. Much evidently remains to be learned in these connections. The simple fact that definitely emerges from our data is that the fronto-occipital head deformation in the newborn of a group is as a rule attended with an increased frequency of development of ear exostoses later in life in that group. The size, forms, and locations of the growths remain apparently unaffected. As to the cause for the increase in frequency, it could be conceived that where artificial head flattening was practiced for a long period of time and favored the development of ear exostoses, the tendency toward these became more and more “alive”, and perhaps also the abnormalities became multiplied through direct heredity in families, thus increasing their frequency in the group, even in the undeformed heads. There is probably a kernel of truth in this, though for the present the idea must remain in the category of speculation. MISCELLANEOUS CAUSES As in other cases where the causation of a condition is obscure, so with ear exostoses, the students of the subject in the course of time have advanced about the whole gamut of theories that could well be made in this connection. Aside from those dealt with in the previ- ous pages may be mentioned the following: INBREEDING Moller-Holst (1932) believes that the pathological ear exostoses, as well as the normal thickening of the tympanic bone such as found in the Eskimo, have arisen through inbreeding (see p. 56). CONSTITUTIONAL DISEASES (OTHER THAN SYPHILIS AND ARTHRITIS ) Heiman (1890) regarded scrofula as favoring a development of ear exostoses, but only indirectly, through its favoring inflammatory con- ditions in the auditory canal. There are a few other weak references 74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 to such a possible connection in the earlier literature. Alexander (1930, p. 456) believes some of the growths to ‘“ be connected with general constitutional diseases’, but does not specify or go further into the subject. INNERVATION ‘ Toynbee’s (1860) class II of ear exostoses “ showed symptoms indicative of diseases in the cavities containing the expansion of the auditory nerve.” CONNECTION WITH EXOSTOSES OF INNER EAR, OR OF OTHER PARTS OF THE SKELETON A connection of meatal exostoses with those of the inner ear was noted by Toynbee, Gruber, Schmaltz. Virchow (1889) saw that in © two of his cases the skeleton showed also other exostoses, which sug- gested a possible connection. Krakauer (1891) was inclined to believe that there was a connection between meatal exostoses and multiple exostoses of the skull. Ostmann (1894) believed the ear exostoses to be due in part to “a tendency to excessive bone produc- tion, such as manifested through multiple exostoses.” OTOSCLEROSIS Burton (1927) attempts to identify the process leading to ear exostoses with otosclerosis. Bezold (1895, p. 50) states that “ sclerotic processes in the middle ear do not infrequently coexist with the exostoses.” MASTOID Rafin and Rougier (q. by Sabroux, p. 22) cite a case of advanced obliteration of the auditory canal by exostoses, in which the mastoids were more developed than the average; Knapp (1898) looked upon some of the ear exostoses—hecause they are so often posterior—as probably due to primary affections of the mastoid. MASTICATION Burton (1927) raises a point which, curiously, has been neglected by other authors. This is the influence upon the meatus of the ac- tivity of the lower jaw. He believes “ irritation to the aural canal due to mastication to be a contributory factor of aural exostosis.” The excess of the growths in the male he attributes to the same factor. But Moller-Holst (1932, p. 102) says the idea of the effect of masti- cation cannot be sustained. NO. 6 EAR EXOSTOSES—HRDLICKA NI (S| VESTIGIAL Whitney (1895) believed “these exostoses must be considered as formed from remnants of fetal cartilage.” NO PATHOLOGY For Le Double and Lebourg (1903), finally, who examined the ear exostoses in a series of American Indian skulls, the growths were “ evidently not of pathological origin.” GENERAL DISCUSSION Including the records presented in this work, there are few if any pathological conditions of the human skeleton that could command as vast an amount of material as that of ear exostoses, not only clinically, but also, and in much larger measure, racially. This ampleness of material enables the student to see the subject in a much more com- plete and satisfactory manner than has hitherto been possible. Let us survey briefly what the facts, as far as revealed, indicate. UNITY OF COMPLEX About the most important result of the studies is the realization that the subject of ear exostoses, notwithstanding its many variations, represents not a mixture of diverse conditions, but in substance a large unit, a special unit-complex, in the field of human derangements. While there is a possibility that some of the smaller growths from the deep portion of the roof of the ear, and now and then perhaps an ossification in a polyp, or a hypertrophy following a serious injury, may be formations apart, the great bulk of exostoses in the external meatus constitutes a single human pathological complex-entity. Re- gardless of any secondary subdivisions of the growths, some of which may be useful, the overwhelming testimony of the evidence is that there is involved but one process—a realization that should facilitate the eventual comprehension of its causes. RESTRICTION TO MAN According to all indications the affliction of ear exostoses is purely human—no such growth has ever been observed in the anthropoid apes, any other primates, or any other living forms provided with an external bony meatus. It is apparently one of the penalties of the human estate. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Ni GENERALIZATION IN MAN Aside from the fact that ear exostoses are limited to the human family, the next major realization is that of the generalization of the tendency to these growths over many if not all the human races. So far the formations have been found in all branches of the human family from which large cranial collections are available. They have not yet been reported in a Negro skull, in that of a Chinese, nor in those of a few other groups, but the numbers of specimens in no one of these cases has been adequate. When sufficient material is available, there are indications that the growths will be found ab- sent in no human aggregate. VARIED FREQUENCY Though a tendency toward ear exostoses is probably a pan-human condition, the incidence of the growths varies greatly in different races and groups, and even in different localities. This may be due to dif- ferences in the tendency, or to the quantity or effectiveness in a given group or locality of the exciting causes; or both these factors to- gether. The most afflicted, in the order named, are some of the American Indians, the Polynesians, and probably some groups of the Whites ; the least affected are the African Negro, possibly the Chinese, and the Eskimo. There is no racial concordance in this and no plain significance, except perhaps as to the African Negro, who in general represents one of the least advanced groups, so far as the brain and head are concerned. The very marked difference in the incidence of ear exostoses among the American Indians and probably also among the Whites indicates that in some groups and localities the agencies that master the normal status have been further weakened, or that conditions favoring the production of the growths are more potent or common. The term “racial inclination ” must be understood in this manner. GENERAL SIMILARITY A survey of large and varied series of osteological material reveals : 1, that in general, ear exostoses display, regardless of race or place, essential similarity; and 2, that there is often distinguishable more or less of group or place peculiarities. Thus in one tribe or locality large growth may be frequent; in another, equally involved, they may be scarce. There are also group differences in the exostoses according to their location, in their predominant forms, and in other respects. No. 6 EAR EXOSTOSES—HRDLICKA V4 It may be said that each more important racial or geographical unit of people will present its own characteristic picture of ear exostoses, though the fundamentals remain much the same. The differences may be of but little import, but now and then are rather striking. All char- acteristic group differences in these respects must necessarily be con- nected, on one hand, with the neuro-vascular background of the growths, and on the other hand, with the status of the normal bony structures involved. NATURE OF THE PROCESS The syndrome of ear exostosis cannot be regarded as a “ disease ”’. Although secondarily it may become pathological and incidentally even fatal, yet in general the outgrowths are quite innocuous, and mostly not even known of, until they grow so large as to become an obstruction or lead to the development of inflammatory conditions. There seems to be no possibility that any bacterial agency is involved in their production, and they are in no sense malignant. If we exclude all microorganisms and all malignancy, then there remain, it would seem, but two classes of possible causes, the first comprising such “ poisons” in the lymph and blood as would be capa- ble of arousing bone tissue to localized bone overgrowths and out- growths, and the other consisting of derangements of the trophic nerve control of the parts concerned. “ Poisonous’, i. e., harmful, substances circulating in the body liquids are certainly present in many individuals, particularly in the later years of life, when the liver, kidneys, and other organs no longer suffice for their neutralization and elimination, and particularly in various pathological conditions and diseases during which poisonous products come into the blood and lymph from pathogenic bacteria. Granted a “ predisposition’ to outgrowths in the tympanic bone and the squamous part of the bony meatus, some such poisons as those just mentioned could readily be conceived as the initiators of the exostoses. What many otologists suspected in connection with syphilis, gout, and other constitutional troubles as causes of ear ex- ostoses, was not really these conditions themselves as much as their poisonous products. Mineral as well as organic poisons, such as lead, mercury, alcohol, etc., may also enter the system in other ways. That some such substances may be capable of inciting abnormal osteogene- sis where the ground for this is favorable may provisionally be ad- mitted. That any special one or ones can excite the bony growths in the ears, would need an unequivocal demonstration through experi- ment. That any and all are generally powerless to produce such bony 6 78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 outgrowths is shown by the very many whose blood and lymph are known to carry such poisons for years, without any effect on the ears. Moreover, such poisons, when present, reach all parts of the osseous system, yet produce nothing resembling the hyperostoses and exostoses of the ear. The case for the internal poisons therefore is not a strong one. There remains the factor termed conveniently “ predisposition ”. This predisposition must be something limited to the external meatus, for barring rare exceptions, no such bony growths appear elsewhere on the skeleton. But what is the “ predisposition”? It can hardly be anything in the mechanics or the structure of the bone—if it were, there would probably be other examples of such a condition and its results elsewhere. There is no evidence that the tympanic bone is unfinished or proliferating in spots. Yet the fault, essentially, can only be with the meatus, or with what controls its structure and being. If the condition of ear exostoses is, as it appears to be, of the nature of an osteogenic derangement, then it would be reasonable to regard it as the result of a disturbed or weakened trophic control of the parts affected. The development of every part of the body is under a very definite and heredity-bound neuro-vascular control. In the apes and lower forms such control of the external bony meatus is evidently thoroughly established and fully adequate. In man this control ap- pears to be disturbed and no longer wholly sufficient; and with, or even without, sufficient exciting causes abnormal bony growths in the ears are the result. Such a disturbance or weakening could possibly be an accompaniment of the unprecedentedly great and rapid evolution of the human head. Such radical change must have disturbed pre- existing trophic controls, and a full reaccommodation has not yet been reached, at least not in most of the human groups. Another, though related, way would be to look upon the deranged neuro-vascular control of the external aural canal as an expression of degeneration. Degeneration may be defined as a generalized, progres- sive groupal insufficiency of the organism to sustain the developmental level reached by an organ. Such insufficiency can again only be based on inadequacy of the trophic nerve centers. It differs diametrically from the inadequacy of accommodation in that it tends to augment with time, whereas the latter tends to diminish, unless the evolutionary changes that caused it keep on advancing. Can the osteogenic disturbance which results in ear exostoses be regarded as a process referable to a pan-human greater or lesser de- generative condition of the tympanic bone or the external meatus? NO. 6 EAR EXOSTOSES—HRDLICKA 79 Normal, that is, primarily devolutionary and not pathological, or- ganic degeneration follows, as a rule, the reduced use of an organ. Examples of it may be observed in human teeth, appendix, the fifth toe, and other parts that in the course of time have become less active and important than formerly. The changes comprise diminution in size, loss of regularity of shape, diminished vital resistance, and a tendency toward elimination of the part. Has the human ear, or at least the external auditory canal, become less useful and necessary to man than it has been in the lower primates and other animals? The changes here, too, aside from the outgrowths, include diminutions in size (lumen), distortions in shape, and occasionally a more or less complete elimination.” ° Is the underlying process of ear exostoses, then, a degeneration, or merely a disturbed accommodation—a slowly regressive, or but a somewhat inadequate central control of the neuro-vascular system of the parts involved, with the bony abnormalities as secondary mani- festations? It may be best not to attempt any answer to this question before the rest of the available facts on the subject can be considered. ANTIQUITY It is not known, and will probably never be determined, when in the existence of man the abnormality of ear exostoses made its first appearance. No case of the growths has as yet been reported in early (geologically ancient) man, and none even from the Neolithic period, though that does not necessarily mean that they were absent. The oldest skulls in which ear exostoses have been encountered so far are the seven skulls with such outgrowths found by me in the Egyptians of the beginnings of the XII Dynasty, or close to 4,000 years ago. These specimens came from the deep rock-tombs at Lisht and belonged to the higher classes of the people. The four next oldest specimens with ear exostoses are the skull of an Egyptian (date?) reported by Ostmann (1894), that of an early Christian Nubian Egyptian mentioned by Wood Jones (1910), and the two Egyptian crania found by me in the collection from the Kharga Oasis. Possibly quite as old, or nearly so, are some of the pre- historic American Indian skulls with the growths. Most of the Ameri- can specimens of such a nature are pre-Columbian; others are post- Columbian but of a period before there was any appreciable influence by the white man. The Hawaiian and New Zealand crania of our ™ See the writer's “ Seven prehistoric American skulls with complete absence of the external auditory meatus.” Amer. Journ, Phys. Anthrop., vol. 17, no. 3, 1933. 8o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 series are probably all pre-White. The European and American Whites recorded in this connection are all recent, as are also the Chinese, the African Negro, and the Melanesians. Thus the time in which ear exostoses are known to have existed extends back about 2,000 years before the Christian era. But the frequency of the condition in the prehistoric American skulls is a sure index that the beginnings of the affection are much older—which is about all that can safely be said on the subject for the present. An even more important question than that of antiquity is that of progression. Have ear exostoses, or have they not, been becoming, however slowly, more frequent? A reliable answer to this query would go far toward the solution of the problem as to whether or not the process is of a degenerative nature; but such an answer is not yet possible. AGE Far exostoses, the data have shown, are encountered from child- hood to old age, but the maximum frequency of their development ranges from adolescence to middle age. With no known exception, they begin to form only after the parts they involve have reached full development, and this development, as seen in the affected skulls, barring an occasional diminution in the lumen of the meatus has as a rule been quite normal. This regular normalcy of structure and the delayed manifestation of the growths speak against the basic cause of the abnormalities being of a degenerative nature. SEX Ear exostoses are decidedly more frequent in males than they are in females. This phenomenon, moreover, is common to all races, all human subdivisions. What is the meaning of this marked and gen- eralized sex difference ? It would be easy to draw heredity into the picture and speculate on its sex dominance and other effects, especially since there is con- clusive evidence that hereditary transmission often does have a part in the spread of these growths. But this method of thinking, with our present knowledge, would lead only to a maze of uncertainties. Yet heredity in the wider sense is involved in the problem, The pan-human extension of that something in the system that predisposes to ear exostoses can only be sustained as time goes on through he- redity ; and this generalized deeper heredity apparently finds its effec- tiveness everywhere enhanced in the males. The females on the whole No. 6 EAR EXOSTOSES—HRDLICKA 81 are more infantile, less differentiated, organically more conservative ; the males more advanced morphologically, less set, more variable. For these reasons the effects of any evolutionary belatedness in ac- commodation of parts could confidently be expected to show more in the males. This seems likely to be the explanation of the sex differ- ence in the incidence of the abnormal conditions under consideration. There may be assumed, in general, a somewhat greater control of the normal conditions of the tympanic bone and the external meatus as a whole in the females. An interesting fact in this connection is that according to pro- . longed observations by otologists, ear troubles and diseases in general are more common in the males than in the females. There is however one notable exception and that is atresia or congenital defect of the external meatus; this serious agenetic defect is considerably more common in the females, particularly in certain human races.” SOCIAL STATUS A number of authors, it has been seen, have voiced the opinion that, among the Whites at least, ear exostoses occur predominantly in the well-to-do classes. If this is so—and there is no voice to the contrary—the question would arise, what is there in the wealthier class that favors the abnormalities? Here is a promising line of inquiry, but the foundations for it need to be strengthened. All that may be said now is that being “ well- to-do’ means, on one hand, usually more leisure and what this carries with it, on the other hand, a greater preservation of the weaker—which, together, may conceivably lead in time to some weakenings or derange- ments of the basic neuro-vascular controls of different parts of the organism. SIDE In materially over one-half of the cases of ear exostoses, on the average, the affliction is bilateral and frequently more or less sym- metrical. This points strongly to the conclusion that the basic cause of the growths lies centrally in the nervous system. That not all cases are bilateral is due probably to developmental differences of the bones of the meatus in the two ears. It is well known that no two bones are exactly equal, macroscopically and microscopically, on the two sides. In the unilateral cases, there is perceptible a general tendency for the exostoses to be more frequent in the left ear. In clinical experience “See the writer’s report on the condition, Amer. Journ. Phys. Anthrop., vol. 17, no. 3, 1933. 82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 with ear troubles as a whole, according to extensive statistics, “ the left ear becomes diseased more often than the right ”, in the proportion of about 5 to 4 (Burkner, 1883, p. 103). The significance of this eludes us for the present.” In this connection it must not be forgotten that not a few one-side cases tend with time to become bilateral. CONSTITUTIONAL DISEASES, LOCAL PATHOLOGICAL CONDITIONS, TRAUMATISMS, IRRITATIONS None of these conditions is associated with ear exostoses, except on more or less infrequent occasions. Examinations of osseous ma- terials show that in cases where gout, arthritis, or syphilis have run unchecked to their limits, there are generally no ear exostoses. These diseases, as well as most of the local irritative conditions affecting the external meatus, can occupy no more than a secondary place in the etiology of the abnormal growths. They can probably all be accepted as occasional exciting, but not as the original, causes, except perhaps in the case of certain traumatisms of the bones composing the meatus. The real, the predisposing, cause is different: it is deeper and is gen- eralized over most if not all human groups. THE GENERAL CAUSE On the basis of the extensive materials reported in this work and after all preceding considerations of the etiology of ear exostoses, both by other authors and the writer, it seems possible now to approach certain generalizations that hitherto were impossible. Let us fix our minds once more on the essential facts of the subject under consideration. These are: an apparently complete absence of the process in animals, including the primates; a generalized predis- position to it in recent and present man; the absence or rarity of developmental defects; the predominance of the manifestations from the post-pubertal to presenile time of life ; the excess of involvement in the males ; the marked tendency of the growths to bilaterality and even symmetry ; and their ascertained occasional direct inheritability. All this, as seen in previous discussion, points to the conclusions that ear exostoses constitute a special complex or entity belonging not in the field of diseases but in that of abnormalities, and that they must be directly connected with neuro-vascular derangements, which “ Here again, curiously, the serious defect of ear atresia forms an exception, being considerably more common, especially in the American Indian, on the right side. No. 6 EAR EXOSTOSES—HRDLICKA 83 may be excited locally by different agencies, but which have their seat in the trophic nervous centers that control the bony structures of the external auditory canal. There remains to be considered the general cause of the neurocentric derangement and insufficiency. It was seen that two possible agencies suggested themselves in this connection. One of these is degeneracy, the other incomplete or disturbed evolutionary adjustment. The disgenic character of the ear growths would seem to point to a degenerative cause affecting the aural apparatus—they tend to lead to loss of hearing, local troubles, and indirectly, in rare cases, even to death. Another item that might conceivably support the view of a degenerative cause of ear exostoses is the fact, long known in otology (see Burkner, 1883, p. 103), that “‘ the disposition to ear troubles (in general) rises from birth to the goth year and thence diminishes with age”, which might be viewed as evidence of a degenerative condition regarding the ear and which bears a close relation to the age incidence of the ear growths. The much greater prevalence of ear exostoses in the males than in the females, in all the human groups known to show the abnormalities, might be viewed as a further support for the idea of a basic degenerative cause, for the males as the less stable or. conservative sex might be expected to make just such a showing. But there are serious difficulties in the way of the hypothesis of degeneration. The most potent are: There is no evidence that the human ear is falling into disuse and hence would be starting on the way to degeneration and restriction or elimination ; ear exostoses affect not the organ of hearing proper, but the relatively much less important bony passage that leads to the hearing apparatus itself ; the exostoses as a rule are not accompanied by congenital defects in the parts in- volved; they are associated with few and followed by very few, if any, changes that could possibly be looked upon as of degenerative nature of either the ear as a whole or of the meatus; once successfully removed by the surgeon they generally leave a sound ear with normal function and but rarely recur; the process of the production of these growths is in substance an excess of production rather than any ex- pression of structural weakness or defective vitality such as usually result from degenerative causes ; and although they may be transmitted to the progeny, the ear exostoses recur as such and not, so far as known, in the form of any congenital defects of the meatus. It appears from the above that there is more that speaks against than for a degenerative cause in the background of ear exostoses. There remains to be considered the central neurotrophic maladjust- ment. It is plain that all structures in the body must be formed and 84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 maintained under the direction of special nervous centers, which as yet are known but very imperfectly but the existence and regulatory function of which are necessary, and which are called the trophic cen- ters. They are highly important parts of the central government of each individual organism. They act upon the blood and lymph supply and doubtless also in other effective ways. The tissues themselves can have no architectonic individuality; they are but so much living material from which, under the strongly hereditarily fixed specific influence of the nervous centers, there are built different structures. Nor does the function of these centers end with the finish of the con- struction, just as the function of the government of a city cannot end with the completion of its streets and houses. There is a perpetual guardianship under the continued power, of heredity, and there are perpetual changes, for heredity is not fully discharged or satisfied with the completion of the structures—it is a lifetime factor. And here, I feel, lies the clue to the problem that confronts us in our study of ear exostoses. It may safely be regarded as axiomatic that as long as both the heredity and the nervous apparatus that subserves it in connection with any given part of the organism are normal, that part will, unless in some way injured, remain normal. If in the absence of a chemical, mechanical, or bacterial injury a part behaves abnormally, it is an un- questionable proof that at that point and to that extent there is either a weakening or derangement of the hereditary control, or of its proper transmission. The something we call heredity must not be conceived as any special power acting within and upon an organism—it can in the end consist only of a specific organization within and between the molecules of the cells of the nervous centers. Such organization is as a rule deeply fixed and not readily influenced. It may however be affected by deep-acting causes. Just what such causes are and how they may act, except in case of destructive bacterial or other poisons, is still but little known. What from the above discussion can help us toward an understand- ing of the basic cause of ear exostoses? Can it be a weakened or temporarily disturbed heredity, or does the cause lie in a faulty or temporarily disturbed mechanism of nervous transmission of the hereditary control ? Both the above conditions are possible, but the wide prevalence of ear exostoses in the human family, and their manifestations as to age, sex, and side, seem to speak against derangements of nerve transmission. No. 6 EAR EXOSTOSES—HRDLICKA 85 This leaves as the most probable basic cause of ear exostoses a weakening or derangement of the normal hereditary control of the tympanic bone and parts adjacent. With this conclusion we must rest for the present, for while it was possible to proceed thus far by the use of logic, a further delving into the subject, until there have ac- cumulated additional reliable data, would entail pure speculation. Only one thought, already touched upon, may here be permissible, but that must not be taken for an assertion. It concerns man’s astound- ingly rapid evolution and changes—geologically and morphologically. Within some 500,000 years man’s progress, especially as concerns his brain and head, has far outstripped that of all the rest of creation. This rapid progress and differentiation, with a spread to all regions and exposure to a multitude of new factors, has prevented in many respects a full adjustment of all parts, a full harmonization and sta- bility in all regions. There is a possibility that the central trophic con- trol of the external meatal region, in the greatly enlarged, altered and still altering skull, has not regained the full life-long adequacy that it possessed before. This would mark the abnormality under con- sideration as an incidental condition, one that might disappear in the natural course of events, if further skull changes affecting the part stopped and if direct inheritance of the abnormality did not meanwhile become rooted. Should this conception prove to be true, then the process of ear exostoses could be defined as that of irregular out- growths of bone in the external bony auditory canal and principally from its tympanic part, due primarily to evolutionarily-weakened normal neuro-vascular control of the parts. One other item demands a brief consideration in these connections. It is that of a possible disturbing, and hence causative, influence of the sex hormones. As seen, ear exostoses develop most frequently during the period of major sexual activity. Also they are more common in the male, in whom the sex activity is more intense. This attractive “clue ”’, however, proves disappointing. Perhaps no race, particularly as regards the males, is more active sexually than the Negro—yet ear exostoses are very rare if not absent. There are so many such incongruities, racial and individual, that this promising lead must be abandoned. SUMMARY Far exostoses are neither a constitutional, nor infectious, nor malig- nant disease. They do not even properly deserve the name of disease, being but secondarily pathological and that mainly through obstruction or pressure. They are abnormalities rather than a disease. 86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Ear exostoses are localized hyperplasias, or outgrowths, from es- sentially the tympanic part, but occasionally also from the squamous portion, of the external bony meatus. They arise generally from what were the free upper ends of the tympanic ring. Their development belongs chiefly to the later adoles- cent period and the earlier half of the adult period. A “ predisposition ” to ear exostoses is now probably universal in man, but in some races or groups the formation of the abnormalities, owing perhaps to direct hereditary effects, is much more frequent than in others. Males are considerably more subject to the growths than females, the well-to-do (in Whites at least) more than the poor. The affection is most frequently bilateral, and where one-sided, it occurs somewhat more commonly in the left than in the right ear. Structurally, the growths range from cancellous to compact, with- out any definite segregation. Though macroscopically the bone is often more or less aberrant, its elements are normal and remain viable. There is never any breaking down or necrosis, nor a com- plete calcification. The causes are systemic or “ predisposing”, and exciting. The paramount systemic cause appears to be a deranged neuro-vascular control of the parts involved, chiefly the tympanic bone, during what corresponds to the sexually more active part of life; but on critical consideration it becomes apparent that no connection of the exostoses with sex activity can be ascertained. 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Eine Exostose im Porus auditorius externus. Arch. Path, Anat. etc., vol. 65, pp. 14-16. No. 6 EAR EXOSTOSES—HRDLICKA gl HARTMANN, A. 1893. Discussion. Berlin. Klin. Wochenschr., vol. 30, p. 636. 1896. Uber Hyperostose des ausseren Gehorganges. Zeitschr. Ohrenheilk., vol. 30, pp. 48-52. Translated by J. A. Spalding, Arch. Otol., vol. 26, pp. 43-46, 1897. Hause, R. 1894. Beitrage zur Wiirdigung der Hyperostose des Felsenbeines. Arch. Ohrenheilk., vol. 37, pp. 161-108. HayMann, D. 1918. Zur Kenntnis der Knochengeschwutlste des Warzenfortsatzes. Wies- baden. HECKE 1893. Operative Entfernung einer Exostose des rechten Gehorganges. Deutsche Med. Wochenschr., vol. 19, p. 552. HEDINGER, A. 1879. Uber Exostose im ausseren Gehorgange und ihre Behandlung. Zeit- schr. Ohrenheilk., vol. 8, p. 379. Translated in Arch. Otol., vol. 9, p. 3, 1880. 1881. Uber eine eigentiimliche Exostose im Ohr. Zeitschr. Ohrenheilk. und Krankh. Luftw., vol. 10, p. 49. Translated in Arch. Otol., vol. 10, PP. 47-54. HEIMAN, T. H. 1890. A case of operation for exostosis of the external auditory meatus. Arch. Otol., nos. 2-3, pp. 144-I5I. HetrricuH, C. H. 1895. Exostoses of auditory canal; operations with dental engine. Homeop. Eye, Ear and Throat Journ., vol. 1, pp. 39-43. HENKE, F., and Lusarscu, O. 1926. Handbuch der speziellen pathologischen Anatomie und Histologie. HoFMANN, LOoTAR 1926. Zur Anatomie des Primaten-Schlafenbeines und seiner pneumatischen Raume unter Berticksichtigung des menschlichen Schlafenbeines. Monatsschr. Ohrenheilk., vol. 60, pp. 921-049. Hovet., T. M. 1894. Removal of an exostosis of the auditory meatus by combined drilling and traction. British Med. Journ., vol. 1, p. 1302. HrouicKa, ALES 1911. Some results of recent anthropological exploration in Peru. Smith- sonian Misc. Coll., vol. 56, no. 16, 16 pp., 4 pls. 1914. Anthropological work in Peru in 1913. With notes on the pathology of the ancient Peruvians. Smithsonian Misc. Coll., vol. 61, no. 18, 69 pp., 26 pls. IrarD, J. E. M. G. 1821. Traité des maladies de l’oreille et de l’audition. Paris. Jackson, G. ; 1909. The etiology of exostoses of the external auditory meatus. British Med. Journ., vol. 2, 1137-1138. g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 JACQUEMART, ALGER 1885. Observation d’une exostose du conduit auditif externe; opération et guérison. Congr. Intern. Otol. Compt.-Rend. Bale, vol. 3, pp. 329- 333; also in Alger Méd., vol. 12, pp. 368-373, 1884. 1889. Communication a propos de deux cas d’exostoses du conduit auditif externe. Rev. Laryng., etc., vol. 9, pp. 189-195 and 218-227. Jones, F. Woop 1910. General pathology. Arch. Surv. Nubia, vol. 2: Report on the human remains (by G. Elliot Smith and F. Wood Jones). Cairo. KAREWSKI 1892. Multiple Exostosen am Schadel und Gesicht mit halbseitiger Gesichts- atrophie. Deutsche Med. Wochenschr., no. 3, p. 57. KAUFMANN, E. 1922. Lehrbuch der speziellen pathologischen Anatomie. 7th and 8th eds. Berlin and Leipzig. Ketson, W. H. 1900. 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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 93), INO== 6, Peat THREE PRE-COLUMBIAN SKULLS FROM PERU, EACH WITH ALARGE POS- TERIOR EAR EXOSTOSIS, WITH VARIOUS BASE ATTACHMENTS SMITHSONIAN MISCELLANEOUS COLLECTIONS Wolbe Cisis INOS a lelbs 2 r ~, A Le & _ Poe 2, ~~ —— a ied THREE PRE-COLUMBIAN SKULLS FROM PERU, WITH IRREGULAR POS- TERIOR AND ANTERIOR (OR A-S) EAR EXOSTOSES SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR Sr INO Ogn eens =~ ay THREE PROBABLY PRE-COLUMBIAN INDIAN SKULLS, UPPER FROM VIR- GINIA, LOWER TWO PERU, WITH DOUBLE (POSTERIOR AND ANTERIOR) EAR EXOSTOSES, OF VARIOUS SHAPES SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE OS NOt iG ele, TWO PREHISTORIC SKULLS FROM PERU WITH EAR EXOSTOSES UPPER: A VERY LARGE POSTERIOR (OR P-1) WITH A SMALL ANTERO- SUPERIOR GROWTH LOWER: TWO LARGE GROWTHS, POSTERIOR AND ANTERIOR, WITH SEC- ONDARY SMALL GROWTHS ANTERIORLY SMITHSONIAN MISCELLANEOUS COLLECTIONS WOES sis INOS G5 tebe Bee Ay ie x ie UPPER: PRE-COLUMBIAN INDIAN SKULL, PERU, WITH TRIPLE EAR EXOS- TOSIS LOWER: THE SAME, WITH A 5 MM-HIGH EXOSTOSIS OUTSIDE ABOVE THE MEATUS, OF SEPARATE ORIGIN AND SIGNIFICANCE. SMALL INTRA- MEATAL EXOSTOSIS IN EACH EAR INDEX OF AUTHORS Alderton, H. A., 18, 60 Alexander, G., 10, 14, 17, 18, 20, 23, 24, 44, 45, 49, 50, 53, 54, 55, 56, 58, 59, 64, 70, 74 Autenrieth, J. H. F., 1, 18, 46 Ayres, S. C:, 18, 34, 40 2S 7 Dr. Ls. of Hamburg,” 67 Bachauer, J., 9, 14, 17, 49 Batlenses, W844. *s2, 55. 60, Gt, 62, 63, 67 Bauer, J., 44 IBEZOld he 7 Tan 2Gn 2Ane27) GO aTe (43) 50, 55, 57, 61, 63, 66, 74 Blakes a Jp s416, os 13) 145 22,130,321, 34, 35, 39, 49, 55, 56, 66 Bonnafont, J. P., 2, 18, 27 Boyer, 55, 56 Braunberger, E., 17, 24, 27, 30, 41, 48, 55 Brindel, 23 Burkhardt-Merian, A., 17 Btrkner, K., 17, 82, 83 Burton, F. A., 9, 10, 14, 20, 34, 49, 52, 54, 55, 64, 74 Carr, LL; 4, 14 Cassells, J. P., 18, 42, 44, 66 Cheatle, A. H., 18 Cocks, D. C., 46, 66 Cook, 66 Dahlstrom, W.., I, 34, 37, 47, 66 Dalby, W. B., 18, 40, 41, 50, 59, 62 Darnach, 54 Davis, J: B.; 3, 14; 22 WelstanchesG. (Gals)h) 175 18, 27. 30; ai. 34, 40, 46, 50, 53, 55, 59, 61, 62 De Rossi, E., 2 Désarenes (see Garrigou-Désarénes ) Dorsey, G., 7, 14 Duplay, 61 Dupuytren, 50 Edward, 24 Erhardt, J., 24, 61 Eulenstein, 46 Perreri, (G:, 18; 23,24, 30, 34, 44, 47, 49 Eield: G: Pe 16; 234 53; 63,65 Flower, W. H., 4, 14, 22 Fournier, 61 Fiirst, 51 Garrigou-Désarénes, 18, 23, 49 Goldstein, M. A., 18, 34, 49, 50, 63, 69 Gray, A. A., 24, 37, 44, 47, 54, 55, 63 Green, Js OF We, 23, 51,63 Geuher, J) 17152, 61.74 Hartmann, A; 5, 13; 18, 34, 42, 44, 52, 55, 56 Hedinger, A., 17, 18, 46, 48, 49, 59, 61, 62, 66 Feiman, IF. EL, 18; 51, 50; 62, 65, 67, 73 Hooton, E. A., 71 Hovyelle ia Mie 18 Jablonowski, 8 Jackson, G., 13, 14, 18, 43, 62, 65 Jacquemart, A., 18, 34, 50, 61, 62, 66, 67 Jones, F. Wood, 79 Karewski, 23 Kaufmann, E., 44, 45 Kerrison, Po D3, 34 Kessel, ©. 'G., a, 13; 18) 23, 24; 27, 34; 39, 40, 41, 42, 44, 47, 49, 51, 55, 56, 50, 61, 62, 63, 66, 67 Kirchner, W., 55, 56 Klotz, 66 Knapp, 74 Korner, (@:,18; 13; 245-25, 27, 30, 38, 34, 40, 41, 43, 51, 55, 56, 57, 63, 65, 67 Krakauer, 23, 61, 74 Kramer, W., 2 Kuhn, 46, 62 Lake, R., 18, 24, 43, 49, 52, 65 ILEKiy, dos 1s, Ge, OS Lebourg, 8, 14, 40, 51, 75 Le Double, A. F., 8, 14, 49, 50, 51, 75 Lucae, A., 17 Manasse, P., 44 Martin, 13 Marx, H., 24, 34, 44, 45, 47, 48 Masini, O., 18, 50, 53, 55, 56, 62 99 TOO Mathewson, A., 18 Mayer, O., 48, 64 McKenzie, D., 44, 63, 65 Meniere, 61 Meyer, A. B., 8 Michailowsk1j, S., 62, 63 Moller-Holst, G., 10, 13, 14, 17, 20, 35, 37) 40, 41, 44, 45, 47, 50, 51, 54 56, 57, 58, 59, 64, 65, 67, 70, 73, 74 Moodie, R. L., 10, 20, 57 Moore, W. O., 8, 18, 55, 56, 65 Moos, S., 34, 48, 51, 62 Nélaton, A., 46, 53 Newark, 17 Noltenius, 66 Noquet, 18, 41, 60, 61 Norrell, 24 Oetteking, B., 10, 14, 20, 22, 34, 44, 47, 70 Ostmatn, Ps 5, 6, 13, 14, 22, 34, 44 53, 69, 74, 79 Patterson, 66 Politzer, “As, 5, 03) 18, 47; 40,49; 525 55, 60, 61, 62, 63, 66, 67 Pritchard, U., 42, 60, 62, 66 Rafin, 74 Ranke, J., 8, 9, 14, 48 Rat Wee 50,102 Reuchard, 2< Roosa, D. B. St. J., 1, 18, 47, 48, 50, 59, 61, 62, 63 Rougier, 74 Russell, F., 8, 14, 16, 70 Sabroux, Ele) 17,08, 27530) 3in34) 37, 39, 41, 43, 48, 50, 60, 61, 62, 66, 67, 74 22:137;°53,0 00} INDEX Schlomkah ji Ga He i7s 1os27) 30.3" 39, 40, 47, 63, 67 Schmaltz, 74 Schwartze, H., 17, 46, 47, 55, 61 Seligmann, 3, 5, 14, 66 Sexton, 2 Siebenmann, 24, 43, 50, 55, 57, 61 Stein, 44 Steinbrugge, H., 1, 48 Stewart, W. R. H., 18, 41 Studleyai@s Ata is. 1422570 Taylor, S. J., 18 Ten Kate, H.,, 7, 14,60, 70 Tod, H., 18, 23) 47,.57, 60, 65,00 Doynbee, J, 1,/2;3, 13,10, 18; saan 42, 46, 58, 50, 62, 74 Triquet, E. H., 61 Turner, W., 4, 14, 49, 60 Urbantschitsch, V., 18, 41 Velpeaux, 8, 50 Wirchow, IR; 3) 5) 14) 34, 42) A4aos 47, 48, 50, 51, 50, 63, 66, 69, 70, 74 Von Luschan, F., 7, 69 Von Rokitansky, C. F., 1, 2 Von Uroeltsch, 2) 24,273 30, si.es4 39, 41, 42, 48, 57, 59, 61 Wagenhauser, 67 Weber, C. O., 47 Weil, 24 Welcker, H., 3, 14, 18, 22 Wiest: G. Ea 18, 23 Whitney, W. F., 24, 27, 37, 48, 49, 68, 60, 75 Wilde, W. R., 62 Wyman, J., 3, 14, 37 Zschokke, E., 3 Zuckerkandl, Emil, 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 7 Mie Con ShANSEN LIGHT FILTER: ITS ADVANTAGES AND LIMITATIONS (WitH Two PLATEs) BY E. D. McALISTER Division of Radiation and Organisms, Smithsonian Institution (PUBLICATION 3297) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION INPAQIE Yio MOSS The Lord Baltimore Press BALTIMORE, MD., U. 8 A. fib CukiSTiANnsEN LIGHT FILTER: ITS ADVANTAGES AND LIMITATIONS By E. D. McALISTER Division of Radiation and Organisms, Smithsonian Institution (WitH 2 PLATES) INTRODUCTION Since the Christiansen light filter is little known in this country, it is believed that a brief description of the filter and a discussion of its possibilities may be useful. The object of the present paper is threefold: 1, to report an improvement in the construction of the filter, which allows its use in an intense beam of light; 2, to discuss the advantages and limitations of these filters for general usage; and 3, to give some “ practical suggestions ” concerning the construc- tion of these filters. The improvement mentioned has arisen from a need (in our laboratory) for an extensive beam of reasonably mono- chromatic light intense enough to produce an easily measured amount of photosynthesis in a higher plant. The second and third purposes of the paper are to answer numerous inquiries the writer has received during the past year. REVIEW OF LITERATURE In 1884 C. Christiansen discovered that a mass of glass particles immersed in a liquid transmitted freely that color for which the liquid and glass particles had the same refractive index. He pointed out in two papers (1884, 1885) that any desired color could be obtained and that a color complementary to the one directly transmitted was seen at oblique angles. He also showed that the wave length of the trans- mitted ray decreased rapidly with an increase in temperature. After a paper with comments and improvements by Lord Rayleigh in 1885, the subject lay dormant for nearly 50 years with the exception of a descriptive paragraph in all editions of R. W. Wood’s “ Physical Optics.” In a series of three papers F. Weigert and collaborators (1927, 1929, 1930) show the necessity of accurately controlling the temperature of the filters and the advantage of a refined optical system, and also describe a single filter that transmits red light when at 18° C. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 7 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 and blue light when at 50° C. Konrad von Fragstein, in 1932 and 1933, describes a filter for the near ultraviolet. One filter covers the range from 3000 A to 3700 A by temperature variation. E. Knudsen, in 1934, discusses all the various ways of making these filters and points out the possibility of making a filter of particles of low-dis- persion glass in combination with particles of high-dispersion glass fused together, both having the same index of refraction for the desired wave length. He has made such a filter but gives no details of its performance. DESCRIPTION OF THE FILTER AND DISCUSSION OF LS ACTION In their commonest form these filters are made up of a solid pack of optical glass particles (0.5 to 2 mm in size) in a glass cell, with the spaces between filled with a liquid having the same index of refrac- tion as the glass for the wave length desired. (The present paper is not concerned with the various emulsions and colloidal preparations exhibiting “ Christiansen colors.’ Readers interested in these are re- ferred to Knudsen, 1934.) Figure 1 gives the curves—index of re- fraction plotted against wave length—for a low-dispersion (borosili- cate) crown glass and a suitable liquid—to percent (by volume) car- bon disulphide in benzene at 20° C. (both anhydrous). Remembering the laws of refraction and reflection at an interface, we see that for the wave length where both liquid and glass have the same index of refraction, the filter acts as a solid plate, and the rays of this wave length are transmitted without deviation or reflection loss within the filter. All other rays of shorter and longer wave lengths are deviated and reflected in an amount dependent upon the difference in the in- dices at the interfaces—glass to liquid and liquid to glass. Examining these curves in figure I more closely, we see that they depart from each other more rapidly on the blue side of the crossing than they do on the red side. This is typical of most suitable glasses and liquids. This shows that the filters will have a sharper blue “ cut-off ” than the red. Also a filter made for blue light will transmit purer colors than one made for longer wave lengths. These two characteristics are evident in the curves shown in figures 2 and 3. Obviously, it is desirable to use a glass of the lowest possible dispersion in combination with a liquid having the highest possible dispersion. The refractive index of a liquid changes rapidly with its tempera- ture in comparison with that of the glass. Hence the color transmitted by the filter will vary with its temperature. Thus to maintain a given color, the temperature of the filter must be held constant. For use NO. 7 CHRISTIANSEN LIGHT FILTER—McALISTER * with light of low intensity, such as in visual work, a carefully thermo- stated water bath is sufficient. lor use with intense light, such as direct sunlight, other more direct means of cooling (discussed below) are necessary. Weigert (1929), making use of this temperature co- 1.54000 800 200 1.53000 800 LIQUID ra 600 400 200 1.52000 INDEX OF REFRACTION 800 600 400 200 1.51000 800 4000 5000 6000 7000 WAVE LENGTH Fic. 1—Index of refraction curves of the components of a filter. efficient, constructed an ingenious filter using methyl benzoate with crown glass particles. This filter transmits red light when at 18° C. and blue at 50° C. These filters are not used like the ordinary colored- glass ones. The “undesired ” colors are not absorbed as in the case of colored glass, but are scattered symmetrically in a halo about the center line through 4 the filter. The angular position of a given SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 ‘ “undesired ” color about the axis of the filter depends upon two factors: 1, the difference in the indices of the liquid and glass for that wave length; and 2, the % TRANSMISSION 45 40 35 30 25 20 iS e A hs ins 4500 5000 5500 6000 6500 JOOO 7500 WAVE LENGTH Itc. 2—Transmission curves of a set of five filters at 20° C. number of interfaces through which the beam passes (i.e., particle size and thickness of the filter). Also, since the interfaces are oriented in a random or probability manner, there exists only a “ most proba- NO. 7 CHRISTIANSEN LIGHT FILTER——McALISTER 5 ble’ angle for a given “ undesired ” color and this color is in evidence in varying amounts at all angular positions about the axis of the filter. Some means of intercepting these “ undesired ” wave lengths is neces- sary. The simplest means is to use an optical system consisting of two lenses with the filter placed between them in parallel light. In this case the “ undesired ” wave lengths are cut out by a diaphragm placed at the image of the source of light. It is imperative that the opening in this diaphragm should conform with the shape of the source of light. If a filament is the source, the opening in the diaphragm should be cut to conform to the shape and size of the image of the filament. If this is not done, the maximum purity of color is not attained. Obviously, the purity of the color obtained increases with an increase in focal length or a decrease in numerical aperture of the optical system used. (See, for instance, von Fragstein, 1933, pp. 33 and 34.) Thus it is necessary to measure the transmission of the filter with the particular optical set-up to be used. Another way of using the filter is in parallel light—direct sunlight, for instance—employing a series of diaphragms to intercept the halo of “ undesired” wave lengths. In this case it is necessary to place the filter at a considerable distance from the ob- server, since the purity of color obtained increases with distance from the filter. Owing to the fact that the “ undesired ” colors are not absorbed but are scattered at various angles about its axis, the Christiansen filter cannot be used in any optical system where sharp images are desired. For instance, it cannot be used before the lens of a camera in pho- tography. The only way it could be used in this respect is with its own optical system to illuminate the object (necessarily a small one) to be photographed. Figure 2 shows the transmission characteristics of a set of five filters at 20° C. They are all made of borosilicate crown glass particles (I to 2 mm in size) immersed in mixtures of carbon disulphide and benzene. The blue filter has about 4 percent (by volume) carbon di- sulphide, the red one 20 percent, and the others have percentages between these limits. These filters are 50 mm in diameter, 18 mm thick, and the windows are fused on optical flats. Two of them are shown in plate 1. The transmission curves (fig. 2) were measured with the filters in parallel light between two 20-cm focal length lenses. A double monochromator and vacuum thermocouple were used to make the measurements. This purity of color is obtained only in the image of the filament used as a source. A battery of Io such filters ranging from ultraviolet to infrared, each selecting a spectral region about 150 A wide (at half maximum), 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 was used by Messrs. Abbot, Stebbins, and Aldrich on Mount Wilson in 1934 to measure the distribution of radiation in the spectra of stars at the Conde focus of the 100-inch reflector. The filters were mounted within a constant-temperature box upon a squirrel-cage device, so as to be successively introduced into a collimated beam. The selected ray was brought to focus with a 19-cm focus lens. All this part of the experiment worked well, and owing to the short-focus objective lens 100 55 %| TRANS. | S 90 80 70 60 % TOTAL INTERVAL ENERGY 50 4450 -4700A 5% RELATIVE ENERGY 4700 - 76 O - 4950 83 4950 - 5200 8 5200 - 5450 25 30 5450 - 5700 1.0 5700 - 6000 0.5 20 4400 4600 4800 5000 5200 5400 5600 5800 6000 WAVE LENGTH Fic. 3—Energy transmission curve for a 6-inch diameter filter equipped with vanes. the quality of the atmospheric seeing was immaterial. Unfortunately, the sharp peak of the photoelectric cell sensitivity proved fatal to the success of the observations. Small traces of stray light of bluish color were so disproportionately effective as to mask real values in the ultraviolet and the red. Dr. Abbot hopes to develop a sufficiently sensitive black receiver as a substitute for the photoelectric cell in future stellar work. Figure 3 shows the energy transmitted by a 6-inch diameter filter (described below and detailed in fig. 4), using the parallel rays of CHRISTIANSEN LIGHT FILTER——McALISTER Lead Gasket Glass Rubber Gasket Aluminum SIDE VIEW 9 _ 1) a ee 2 oO § cv 7 ca 3 38 a o aw y = re es = ro) allt /\6 INCHES FACE VIEW Fic. 4.—Diagrams showing detailed construction of a 6-inch filter with vanes. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 direct sunlight. The observations were made 66 feet from the filter with the filter temperature held at 20° C. This purity of color is ob- tained over a 6-inch circular area at that distance. AN IMPROVEMENT THAT PERMITS THE USE OF AN INTENSE BEAM OF LIGHT The necessity of an optical system and the disadvantage of the temperature coefficient have been pointed out in detail in previous publications. However, for any exacting use of the filter where much light energy is used there is still another trouble which proves to be very serious: when a strong beam of light is forced through the filter, some energy is absorbed, and the center reaches a higher temperature than the edges, owing to poor heat conduction. The color transmitted is no longer pure, even when the filter is in a water hath. The writer has finally overcome this difficulty by inserting aluminum vanes through the body of the filter so as to cut off the least amount of light and carry off as much heat as possible. Details of a satisfactory filter equipped with these vanes are shown in figure 4. The body of the filter is cast aluminum, machined as shown, to take the glass windows and the necessary gaskets. No cement is known to the writer that will satisfactorily withstand the benzene and carbon disulphide mixture on the inside and the water on the outside. For this reason the windows were clamped on, as shown, with a soft lead gasket (4 inch thick) underneath the glass. Ridges were machined on the aluminum face to press into the lead gasket and improve the seal. A zig-inch rubber gasket is placed between the glass window and the brass clamping ring. The vanes are of 35-inch aluminum assembled so that their extremities press firmly against the inner wall of the aluminum case, thus providing a path of good heat conduction from the inside of the filter to the surrounding water bath. The holes shown in the vanes allow the cell to be filled with the glass particles after it is assembled. Without these vanes, the center of this filter rose 9° C. above the temperature of the water bath when the rays from a 1,000-watt lamp were concentrated on the filter. With the vanes installed, the temperature at the center of the filter rose only 0.25° C. above that of the water bath under the same conditions. Plate 2 figure 1, is a photograph of this filter in its water bath. The filter is filled with glass particles and a liquid (about 9 percent carbon disulphide in benzene), and gave the transmission curve shown in figure 3 under the conditions previously mentioned. Plate 2, fig- ure 2, isa photograph of a 12 X 14-inch filter (not filled) with its water bath. When in use the water of the bath is thermostated and stirred. NO. 7 CHRISTIANSEN LIGHT FILTER—McALISTER 9 Aluminum was chosen as the metal for the cell and vanes because it is least attacked by the various liquids used. The outside of the alumi- num case must be carefully covered with several coats of waterproof paint. PRACTICAL CONSIDERATIONS IN THE CONSTRUCTION AND USE OF THESE FILTERS The Christiansen filter is little known in this country. For this reason the writer believes it will not be amiss to pass on to those in- terested some practical points concerning the construction of these filters and their uses. In this connection the writer is drawing on the literature cited and his own experience with these filters. The type of cell chosen to hold the components depends upon the use to be made of the filter. For visual work and other uses where only moderate intensities are necessary, a glass cell with parallel win- dows fused on is suitable. If a permanent filter is desired, a small expansion chamber should be provided on the filling “ neck,’ and after filling, the cell should be sealed off above the expansion chamber: ina flame. To do this safely the expansion chamber should be packed in carbon dioxide snow. When high intensities of light are used, such as direct sunlight, the cell needs to be of the type detailed in figure 4. To be sure, a thin glass cell may be used for high intensity work, but the purity of color will be very inferior to that obtained with a metal- cased filter equipped with vanes. The glass particles for the filter should be of the best optical quality obtainable—preferably low-dispersion borosilicate crown glass. Fused quartz is also suitable and of course necessary for ultraviolet work. However, the quartz should be free of bubbles and inclusions, as these lower the transmission of the filter and give it a muddy appearance. In preparing the glass particles, the writer has used the following pro- cedure. If the glass or fused quartz is in large fragments, it is ground up with an iron mortar and pestle until the larger particles are 2 or 3 mm in size. This should be done with a minimum of grinding. A damp towel should be wrapped about the top of the pestle and draped over the top of the mortar to prevent the “dust” from flying. The operator should use some protection over his nostrils to avoid breath- ing the dust. The glass particles are graded by running them through several sizes of sieves. Before using, the particles must be carefully cleaned. This is best accomplished by boiling in chromic acid cleaning solution. The particles are then washed many times in clear water, then in distilled water, and finally dried completely. The particle size found suitable by previous workers and the writer ranges from 0.5 IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 to 2 mm (usually graded closer—i. e., 0.5 to I mm, or I to 2 mm). A cell 15-mm thick using the 0.5 to 1 mm particles gives approximately the same results as a 30-mm cell using the 1 to 2 mm particles. With a given optical system, reducing the particle size or increasing the cell thickness gives a narrower transmission curve with lower percentage transmission at the “ peak.” In an “ideal”’ filter the glass particles would be perfectly homogeneous as to refractive index, and these par- ticles and the liquid surrounding them would be all at exactly the same temperature. It is because these two conditions can never be realized that the percentage transmission at the “ peak ” decreases as the num- ber of interfaces in the filter is increased. In filling the cell it is best to put the liquid in first—enough to fill the cell about half full. The glass particles are then poured in slowly so that air bubbles are not carried down. It is difficult to free the cell of air bubbles after it is packed solid with the glass particles. The liquid or liquids used must be anhydrous and of the highest purity. The mixing of carbon disulphide and benzene—originally suggested by Chris- tiansen in 1884—to obtain a liquid of any desired index of refraction (between that of pure benzene and pure carbon disulphide, of course) has been found very satisfactory by the writer in spite of its relatively high temperature coefficient. Methyl benzoate, used by Weigert (1929, 1930), in combination with crown glass particles makes a remarkably variable filter. Von Fragstein (1932, 1933) uses a mixture of 44 per- cent alcohol (ethyl) and benzene with fused quartz particles for an ultraviolet filter. This filter, with suitable optics, transmits a narrow band of wave lengths in any desired part of the region 3000 A to 3700 A. The wave length of maximum transmission is shifted as desired in this region by temperature variation, just as in Weigert’s methyl benzoate cell. Various optical systems have been described in the literature. Wei- gert’s (1929) autocollimator is of considerable interest as it passes the rays twice through the filter. The writer has shown that the filter can be used successfully without an optical system (other than plane mirrors and diaphragms ) in direct sunlight, or, of course, in any beam of similar parallelism of rays. In using an optical system it is again emphasized that the diaphragm at the image of the source of light must conform in size and shape to this image. Any change in this diaphragm will change the transmission characteristics of the set-up. This is shown clearly by Weigert (1929, fig. 13, p. 159). In the use of the filter for studying the wave-length effect of some photochemical phenomena it is necessary to allow for or take into account the effect of the “ undesired ”’ colors—i. e., those wave lengths ‘ NO. 7 CHRISTIANSEN LIGHT FILTER—McALISTER a shorter and longer than the wave length of maximum transmission. In any case care must be exercised in determining the combined effect of the shape of the transmission curve of the filter, the wave length versus sensitivity curve of the phenomena under investigation, and, in the case where the energy content of the beam from the filter is measured with a photocell, the sensitivity versus wave-length response of the detector. For instance, if the energy in the beam from an ultra- violet filter is measured with a photocell that has its maximum sensi- tivity in the blue, considerable error may come into the final result owing to the long-wave-length “tail”? on the transmission curve of these filters. (See von Fragstein, 1933, fig. 8, p. 33.) The writer believes that these filters will be found of considerable value as a source of monochromatic light for rough visual measure- ments of refractive index, rotation of plane of polarization, etc., be- cause one can set cross hairs on the wave length of maximum trans- mission within + 10 angstroms. With a sealed filter and accurate temperature control this wave length of maximum transmission 1s sharp and reproducible. The large filter shown in plate 2, figure 2, will be used with sunlight to irradiate a growing plant in an experiment to determine the wave- length effect of photosynthesis. At great distance it will yield a trans- mission curve comparable to that shown in figure 3. Two filters are to be used—one to cover the range 4000 A to 6000 A, the other from 5500 A to 8000 A. The wave length of maximum transmission is moved through these ranges by temperature variation. The possibility of substituting a high-dispersion glass for the liquid—i. e., making the filter of a high-dispersion glass flowed around the particles of low-dispersion glass—is interesting. Knudsen (1934) has accomplished this, but gives no details. The resultant filter will have only a very small temperature coefficient, which will considerably enhance its usefulness. The writer has in his possession two suitable glasses, but has not yet had an opportunity to complete the filter. The writer is grateful to A. N. Finn, of the United States Bureau of Standards, who has kindly furnished the borosilicate crown glass, and to L. B. Clark, of the Division of Radiation and Organisms of the Smithsonian Institution, who constructed the glass cells with fused- on optical windows. LITERATURE, CIVED CHRISTIANSEN, C. 1884. Untersuchungen iiber die optischen Eigenshaften von fein vertheilten Korpern. Ann. Phys. Chem., vol. 23, pp. 298-306. 1885. Idem, vol. 24, pp. 439-446. I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 KNUDSEN, E. 1934. Chromatische disperse Zweiphasensysteme unter besonderer Beriick- sichtigung ihrer Verwendung als Lichtfilter. Koll. Zeitschr., vol. 66, pp. 257-266, March. Konn, H., and von Fracstein, K. 1932. Ein Christiansenfilter fiir ultraviolettes Strahlung. Phys. Zeitschr., vol. 33, pp. 929-931. . RAYLEIGH, Lorp 1885. On an improved apparatus for Christiansen’s experiment. Phil. Mag., vol. 20, pp. 358-360. VON FRAGSTEIN, K. 1933. Ein Christiansenfilter fiir ultraviolettes Licht. Ann. Phys., vol. 17, Pp. 22-40. WEIGCERT, F., and Staupe, H. 1927. Uber monochromatische Farbfilter. Zeitschr. Phys. Chem., vol. 130, pp. 607-615. WEIGERT, F., Straube, H., and Etvecarp, E. 1929. Uber monochromatische Lichtfilter. II. Zeitschr. Phys. Chem., ser. B, vol. 2, pp. 149-160. WEIcERT, F., and SHIDEr, J. 1930. Photodichroismus und Photoanisotropie. VII. Ibid., vol. 9, pp. 329- 355. (See pp. 331-334.) Woop, R. W. Christiansen’s experiment. In all editions of Wood’s Physical Optics. ~ee 5! ST14D SSV19D NI SYALTIA HONI-OM_L Eqs] BYE Mey] SG “UWoya' SNOILO311090 SNOANVIISOSIN NVINOSHLIWS HLVG YSLVYM HLIM YSLT1I4 LHOITD NASNVILSIYHD HONI-PLXZIL V CZ HLVG YSLVM HLIM YSLT1IA LHOSOIT NASNVILSIYHD HONI-G VY CL Gad 2) ON) (£6) S1OA SNOILO31100 SNOANVITISOSIN NVINOSHLIWS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 8 fee eho rCALION: OF THE EDRIOASTEROIDEA (WITH ONE PLATE) BY R. S. BASSLER Head Curator, Department of Geology, Wiese National Museum (PUBLICATION 3301) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION APRIL 4, 1935 The Lord Waftimore Press BALTIMORE, MD., U. 8. A. THE CLASSIFICATION OF THE EDRIOASTEROIDEA BYaRe Ss BASSEER Head Curator, Department of Geology, U..S. National Museum (WitH ONE PLATE) The Edrioasteroidea, a group of Paleozoic echinoderms, regarded by specialists either as a distinct class of the Pelmatozoa allied to the cystids or as an aberrant order of the same division, has been the subject of so many detailed observations and changes in nomenclature that much confusion exists in its classification. Many of the genera have inexact limits assigned them, because they are based upon incor- rect illustrations or described from species other than the cited geno- type, or, again, because they are founded upon characters of uncertain value. Lastly, some authors, without regard to. the rules of nomen- clature, changed the generic endings to suit their ideas as to the relationships of the group, Agelacrinites, for example, becoming first Agelacrinus and later Agelacystis. Furthermore, it must be remem- bered that the camera-lucida drawings of years ago resulted in reversed images, the right side becoming the left, which in the edrioasteroids gave rise to a serious error since the direction of the ambulacra, a generic character now believed to be important, thus was reversed. However, most students paid little attention to the extent and direction of curvature of the ambulacra, including in the same genus forms most diverse in these respects. In preparing the present classification the writer first of all proved, at least to himself, from a study of several hundred specimens of several Cincinnatian species, that the amount of imbrication of the interambulacral plates, the width of the peripheral border of plates, and the number of arms or ambulacra may vary even in the same species ; but the plate structure of the ambulacral and oral areas, and the direction and extent of curvature of the ambulacra, remained constant, thus affording good generic characters. In their simplest form the edrioasteroids have a flexible theca or sacklike body composed of numerous more or less polygonal plates having a mouth on the upper surface at the central point of radiation of five straight or curved arms or ambulacra separated by interam- bulacral plates, and an anus with valvular covering occupying part SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 8 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 of an interray with a hydropore located between it and the mouth. These primitive echinoderms, free or attached by a part of the lower surface, developed into parasitic circular flattened forms, into elevated sacklike bodies or into cylindrical objects bearing the ambulacra at the top of a fused mass of plates. Starting at the left of the anal area and proceeding clockwise, the ambulacra are designated as follows: 1, left posterior ; 2, left ; 3, anterior (opposite the anal area) ; 4, right ; 5, right posterior. These numbers are employed in the following descriptions. Through the study of the Springer and Ulrich collections of Edrio- asteroidea in the National Museum, and of material lent by other insti- tutions, the writer has been enabled to review the structure of most of the known genera and species of this group, the generic results of which are offered below. Here, the old genera are briefly redefined and six new generic names are proposed. This paper is introductory to a more complete publication that will include descriptions of additional new species upon which this classification is based. Class EDRIOASTEROIDEA Billings, 1854-58 (Thyroidea Chapman, 1860; Agelacrinoidea S. A. Miller, 1877-1883; Cys- tasteroidea Steinman, 1888; Thecoidea Jaekel, 1895; Cystostellaroidea Steinman, 1904. ) Family AGELACRINITIDAE, new name (Agelacrinidae Jaekel, 1899; Thecocystidae Jaekel, 1899) Theca flexible, composed of thin plates attached temporarily or permanently by the greater part of the aboral surface and with am- bulacra confined to the oral surface. A single row of ambulacral flooring plates overlapping proximally. STROMATOCYSTITES Pompeckj, 1896 (Stromatocystis Bather, 1900) Theca depressed, pentagonally globular in form with oral surface bearing five narrow, straight ambulacra separated by polygonal mosaic interambulacral plates with numerous intervening pores and the under surface completely occupied by smaller, less regular polygonal plates. Anal area slightly elevated, indistinct. Genotype.—S. pentangularis Pompeckj, 1896. Middle Cambrian of Bohemia. S. balticus Jaekel, 1899, S. walcotti and var. minor Schuch- ert, 1919 are other species of the genus. no. 8 EDRIOASTEROIDEA—BASSLER 3 WALCOTTIDISCUS, n. gen. Pentagonal form and general shape of theca as in Stromatocystites, but the oral face bears curved ambulacra, four (1-4) directed to the left and one (5) to the right. Furthermore, the edge of the oral sur- face is composed of a narrow band of very small nodose plates and the under side of larger polygonal plates imbricating. Anal area indistinct. Genotype.—W. typicalis, new species. Middle Cambrian of British Columbia. WALCOTTIDISCUS TYPICALIS, n. sp. late siete. oe Type specimen, free, depressed, globular, subpentagonal in outline, imbedded and somewhat distorted in hard shale. Oral face with five slightly curved ambulacra, four directed to the left and one, the right posterior, to the right, separated by polygonal interambulacral plates very slightly overlapping. Edge of oral side formed by a band con- sisting of five or more rows of very small elongate imbricating plates ; basal side of theca as shown where pushed over oral face, occupied by large elongate polygonal, imbricated plates. Anal area not dis- tinguishable with certainty. Occurrence-—Burgess shale of Middle Cambrian, Burgess Pass near Field, B.\C..(loc..35 k)- Holotype —U.S.N.M. no. 90754. CYSTASTER Hall, 1871 (Thecocystis Jaekel, 1899) Theca an elongate sack with the lower end attached to some foreign object. Ambulacra short and straight, five in number, with plates often rather nodose. Interambulacrals rounded or polygonal, mosaic, very minute (0.25 mm in diameter). Anal pyramid of Io elongated, abruptly raised plates. Genotype.—Hemicystites (Cystaster) granuatus Hall, 1871. Or- dovician (Maysville) of Ohio. CINCINNATIDISCUS, n. gen. (Hemicystites of authors not Hall) Like Cystaster except that the theca is not sacklike but depressed, attached, and that the interambulacral plates are squamose and imbri- cate distinctly, divided into larger central plates and a marginal zone of small nodose ones. Anal pyramid of nodose plates. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Genotype.—Agelacrinus (Hemicystites) stellatus Hall, 1866. Or- dovician (Maysville) of Ohio. Hemicystites carnensis Foerste, 1914, belongs to this genus. CARNEYELLA Foerste, 1916 Theca typically elevated, sacklike, attached by narrow basal part but also may occur as a thin circular expansion. Ambulacra curved, four (1-4) to left and one (5) to the right, with two rows of covering plates elevated and alternating with each other along the middle line. Oral area composed of three plates, one large and two small. Anal area of many small plates radially arranged. Surface of plates orna- mented with minute pits and nodes. Genotype. —Agelacrinus (Lepidodiscus) pileus Hall, 1866. oe vician (Maysville) of Ohio. Carneyella vetusta Foerste, 1914, i related to C. cincinnatiensis, n. sp., next described. Carneyella ene rensis Clark, 1920, is doubtfully referred here. CARNEYELLA CINCINNATIENSIS, n. sp. Plate 1, fig. Io Agelacrinus (Lepidodiscus) cincinnatiensis Hall (not Roemer), Descr. new species fossils Cincinnati, Ohio, p. 214, pl. 2, fig. 7, 1871 (advance sheets, p. 6, 1866) ; 24th Rep. New York State Cab. Nat. Hist., p. 214, pl. 6, fig. 7, 1872. This well-marked species, erroneously referred to Agelacrinus (now lsorophus) cincinnatiensis Roemer, is readily distinguished from that form by its thin, depressed, almost flat theca attached by its entire aboral portion. Furthermore, the ambulacral plates are less elevated, all the plates are rather papillose, and there are only three oral plates. Occurrence.—Maysville division (Corryville beds) of the Ordo- vician, Warren County, Ohio. Holotype-—U.S.N.M. no. 40743. ISOROPHUS Foerste, 1916 Theca depressed, circular, attached by the entire under surface. Ambulacra composed of two rows of plates with an intercalated series often present, more or less curved, four (1-4) to the left and the right posterior (5) to the right. Oral area occupied by numerous small plates. Margin composed of numerous rows of small plates slightly increasing in size inward. Interambulacral plates polygonal, becoming slightly imbricated. Anal area a circle of triangular plates regularly arranged. no. 8 EDRIOASTEROIDEA—BASSLER 5 Genotype—Agelacrinus cincinnatiensis Roemer, 1851. Ordovician (Maysville) of Ohio, etc. Other species of Jsorophus are Agela- crinus austini Foerste, 1914, A. faberi Miller, 1894, A. warrenensis James, 1883, and A. holbrooki James, 1878. ISOROPHUSELLA, n. gen. Like Isorophus but ambulacra I, 2, and 3 directed to the left and 4 and 5 to the right, and the interambulacral plates strongly imbricating. Genotype—Lebetodiscus inconditus Raymond, 1915. Ordovician (Trenton) of Ontario, Canada. ISOROPHUSELLA INCONDITUS (Raymond) Plate 1, fig. 11 Lebetodiscus inconditus Raymond, Ottawa Nat., vol. 29, pl. 1, fig. I, 1915. Theca a thin, slightly convex, circular disk, attached to or resting upon a limestone bed by its entire aboral surface, about 20 mm in diameter, exhibiting on its oral face a broad peripheral band of minute, closely imbricating plates, five ambulacral areas of which the oral por- tion is composed of many small irregular, angular plates, and each ambulacrum has two rows of covering plates alternating with each other and frequently developing additional small accessory plates along the midline between them. Ambulacra short, rather broad, three (1-3) curving to the left and two (4, 5) to the right. Inter- ambulacral areas of larger, narrow, strongly imbricating plates. Anal area rather distinct, consisting of Io elongate triangular smooth plates in one circle meeting at the center but not rising above the general surface. Although very similar to species of [sorophus, this interesting form is readily distinguished by the different arrangement of the ambulacra. Occurrence.—Trenton limestone, Hull, Quebec. Plestotype —U.S.N.M. no. 8.3871, Springer Coll. STREPTASTER Hall, 1872 Theca as in Carneyella, except that the ambulacra all revolve strongly toward the left and are composed of highly elevated plates. Interambulacrals are small, polygonal, mosaic plates, 0.5 to 1.0 mm in diameter. Genotype.—Agelacrinus vorticellatus Hall, 1866. Ordovician (Maysville) of Ohio. S. reversatus Foerste, 1914, and S. septem- brachiatus Miller and Dyer, 1878, are other species of Streptaster. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 LEBETODISCUS Bather, 1908 Theca depressed, circular. Ambulacra curved, five in number, all directed toward the left, with mosaic interambulacral plates. Am- bulacral plate structure apparently as in Carneyella. Anal area well defined in slightly wider interradius. Genotype.—Agelacrinites dicksoni Billings, 1857. Ordovician (Trenton) of Ontario, Canada. L. loriformis Raymond, 1915, is a second described species. FOERSTEDISCUS, n. gen. Like Lebetodiscus, but ambulacra all directed to the right, short, broad, and strongly curved, and interambulacral plates polygonal but overlapping toward the center and highly imbricating around the periphery. Anal area of many small, irregularly arranged plates in slightly wider interradial area. Genotype-—F. grandis, new species. Ordovician (Trenton) of Kentucky. FOERSTEDISCUS GRANDIS, n. sp. Plate 1, fig. 12 Theca a flattened disk, 25 mm in diameter, resting upon or attached to a layer of limestone. Ambulacra five, short, broad, all curved to the right, consisting of two rows of elongate covering plates slightly raised along the midline, and a single row of floor plates. Interambu- lacrals large, imbricating, but flat in the central areas, narrow, more numerous, and piled up on edge along the margin. Anal area large, distinct, occupying a slightly wider space, composed of many small imbricating plates arranged in circles, the smallest in the center. Although resembling Lebetodiscus dicksoni (Billings), the right- handed arrangement of the ambulacra in this present species is a ready means of separation. Occurrence—Curdsville division of the Trenton, near Troy, Wood- ford County, Kentucky. Holotype —U.S.N.M. no. S.3191, Springer Coll. PYRGOCYSTIS Bather, 1915 Oral surface of five broad, straight ambulacra mounted on a high cylindrical turret composed of wide, thin, closely imbricated plates. Genotype—P. sardesoni Bather, 1915. Ordovician (Decorah) of Minnesota. Scalpellum sulcatum, procerum, and cylindricum Aurivil- no. 8 EDRIOASTEROIDEA—BASSLER 7 lius, 1892, Pyrgocystis octogona Richter, 1930, P. grayae Bather, 1915, P. ansticei Bather, 1915, and P. batheri Ruedemann, 1925, are the other species of this genus. HEMICYSTITES Hall, 1852 (Hemicystis Haeckel, 1896) Theca a thin, flattened disk with oral arrangement and pitted plate structure of Carneyella, but ambulacra are short, broad, and straight. Anal pyramid elevated and composed of a circle of triangular plates regularly arranged. Genotype.—H. parasitica Hall, 1852. Silurian (Rochester) of New York. Agelacrinites bellulus, bohemicus, confertus, latiusculus, sim- plex, tener, and velatus of Barrande, 1887, A. rectiradiatus Shideler, 1918, and Agelacrinus billingsi Chapman, 1860, Lebetodiscus youngi Raymond, 1915, and L. chapmani Raymond, 1915, are described spe- cies of Hemicystites. THRESHERODISCUS Foerste, 1914 Agelacrinitidae with branched ambulacral rays having a pronounced trimerous origin. Interambulacrals large, squamose, imbricating in central part and smaller along border. Genotype-—T. ramosus Foerste, 1914. Ordovician (Trenton) of Ontario, Canada. AGELACRINITES Vanuxem, 1842 (Agelacrinus authors; Agelacystis Haeckel, 1895; Haplocystites Roemer, 1852; Haplocystis Bather, 1899) Thin parasitic disks attached by entire aboral surface. Ambulacra five, long, narrow, much curved; two (4, 5) to the right and three (1-3) to the left. Interambulacral plates mosaic and sculptured, al- though slightly imbricating. Periphery formed by several rows of small plates followed by one row of much larger elongate ones. Anal area a circle of triangular, regularly arranged plates. Genotype —A. hamiltonensis Vanuxem, 1842. Devonian (Hamil- ton) of New York. A. ephraemovianus Bogolubov, 1926, A. hanoveri Thomas, 1924, and A. rhenanus Roemer, 1851, from the Devonian, and A. blairi Miller, 1894, A. legrandensis Miller and Gurley, 1894, from the Mississippian are referred to this genus. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 DISCOCYSTIS Gregory, 1897 (Echinodiscus Worthen and Miller, 1883 (not Agassiz) ; Ageladiscus Miller, 1897) Like Agelacrinites except that four ambulacra are curved to left and one, the right posterior (5), to right, and that body is well de- veloped and sacklike with many rows of elongate, narrow marginal plates. Genotype—Echinodiscus optatus Worthen and Miller, 1883=D. (Agelacrinus) kaskaskiensis Hall, 1858. Mississippian of Illinois. Echinodiscus sampsoni Miller, 1891, is another species of this genus. COOPERIDISCUS, n. gen. Body depressed globular, free, entirely covered with plates ; ambu- lacra long, very narrow, much curved, all turning to the right. Inter- ambulacral plates strongly imbricate. Anal pyramid of regular tri- angular plates meeting at a point. Genotype.—Lepidodiscus alleganius Clarke, 1901. Devonian (Che- mung) of New York. LEPIDODISCUS Meek and Worthen, 1868 Like Cooperidiscus but ambulacra 1-4 curve strongly to the left and 5, the right posterior one, to the right. Interambulacral plates small, strongly imbricating. Anal pyramid well developed. Genotype.—Agelacrinites squamosus Meek and Worthen, 1868. Mississippian. Agelacrinites beechert Clarke, 1901, A. buttsi Clarke, 1901, Lepidodiscus lebouri Sladen, 1879, and L. muilleri Sharman and Newton, 1892, are the known species of this genus. ULRICHIDISCUS, n. gen. Body semiglobose with the oral side occupied by five narrow but well-defined, very long ambulacra all strongly curving to the left, with a well-defined anal pyramid of many long triangular plates in one circle, and interambulacral areas composed of polygonal but slightly imbricating plates. Genotype—Agelacrinus pulaskiensis Miller and Gurley, 1894. Mis- sissippian (Chester) of Kentucky. Family EDRIOASTERIDAE Bather Theca flexible, depressed, usually globular, attached by the small central excavated part of the aboral surface; ambulacra strongly No. 8 EDRIOASTEROIDEA—BASSLER 9 curved and passing on to aboral surface; floor plates arranged in two series, one on each side of the ray, and alternating along the median line. EDRIOASTER Billings, 1858 (Cyclaster Billings, 1857, not Cotteau, 1856; Agelacrinites Forbes, 1848, not Vanuxem; Edriocystis Haeckel, 1896; Aesiocystites Miller and Gurley, 1894; Aesiocystis Bather, 1900) Sacklike, flexible theca attached by a small central portion of the aboral surface and with the ambulacra passing from the oral to aboral sides. Interambulacral plates mosaic. Ambulacra strongly curved, four (1-4) to the left and one (5) to the right, with an anal pyramid composed of many small plates, irregularly arranged. Genotype.—Cyclaster bigsbyi Billings, 1857. Ordovician (Trenton) of Canada. Agelacrinites buchianus Forbes, 1848, Edrioaster levis Bather, 1914, E. saratogensis Ruedemann, 1912, and Aesiocystites priscus Miller and Gurley, 1894, are other species of this genus. DINOCYSTIS Bather, 1898 Oral surface convex, with five narrow ambulacra all strongly curved to the left and aboral surface composed of a thin flexible integument of narrow imbricating plates with a small central orifice for attachment. Genotype.—D. barroisi Bather, 1898. Devonian of Belgium. Family CYATHOCYSTIDAE Bather Edrioasteroidea in which the aboral portion consists of a fused solid mass of plates attached permanently to some foreign object. CYATHOCYSTIS Schmidt, 1880 Theca with oral surface much as in Stromatocystites but mounted upon a solid fused mass of stereom and permanently attached at the aboral end. Genotype-—C. plautini Schmidt, 1880. Ordovician (Echinosphe- rites limestone) of Estonia. CYATHOTHECA Jaekel, 1927 Like Cyathocystis, except that the ambulacral areas are apparently very narrow, practically hidden at the surface. Genotype.—C. suecica Jaekel, 1927. Ordovician of Sweden. 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 POSITION UNCERTAIN Family ASTROCYSTITIDAE, new name (Steganoblastidae Bather ) ASTROCYSTITES Whiteaves, 1897 (Steganoblastus Whiteaves, 1897) Regarded by Bather as an edrioasteroid and by Hudson as a blastoid. Family CY CLOCYSTOIDIDAE, Sy Aj Miller, 2889 (order uncertain) Genera Cyclocystoides Billings and Salter, 1858; Narrawayella Foerste, 1920; Savagella Foerste, 1920. EXPLANATION OF PEATE Fic. 1. Walcottidiscus typicalis, new genus and species. The holotype, 3, a crushed theca showing the oral side. Middle Cambrian (Burgess shale) Burgess Pass, near Field, British Columbia. Fics. 2, 3. Carneyella (Agelacrinus) pileus Hall, 1866. View of oral side of theca and lateral view, 2, showing the sacklike form. Maysville (Fairmount formation) : Cincinnati, Ohio. Fic. 4. Cystaster (Hemicystites) granulatus Hall, 1871. Side view, X 2, of the elongate theca. Maysville (Fairmount formation): Cincinnati, Ohio. . Streptaster (Agelacrinus) vorticellatus Hall, 1856. Oral side of theca slightly enlarged, exhibiting the elevated ambulacra all revolving toward the left. Maysville (Fairmount formation): Cincinnati, Ohio. Fic. 6. Cincinnatidiscus (Agelacrinus) stellatus Hall, 1866. Oral side of the depressed, attached theca, with straight arms, 2. Maysville (Fair- mount formation) : Cincinnati, Ohio. Fic. 7. Ulrichidiscus (Agelacrinus) pulaskiensis Miller and Gurley, 1894. View of the semiglobose theca, natural size, showing strong curvature of all ambulacra to the left. Chester (Glen Dean formation): Pulaski County, Ky. Fic. 8. Pyrgocystis sardesoni Bather, 1915. Side view of theca, < 14, showing the solid basal portion of many imbricating plates surmounted by the oral surface. Black River (Decorah) shales: St. Paul, Minn. Fic. 9. Cooperidiscus (Lepidodiscus) alleganius Clark, 1901. Oral side of the free, globular theca, X 14, exhibiting the very narrow ambulacra all curved to the right and the highly imbricated interambulacral plates. Chemung formation: New York. Fic. 10. Carneyella cincinnatiensis, new species. View of the attached, thin cir- cular theca with ambulacral structure of Carneyella, X 4. Maysville (Corryville formation) : Warren County, Ohio. Fic. OL e = no. 8 EDRIOASTEROIDEA—BASSLER Vat Fic. 11. Jsorophusella inconditus (Raymond, 1915). Example, * 4, a depressed circular expansion with the plate structure of /sorophus but with 3 of the ambulacra directed to the left and 2 to the right. Trenton limestone: Hull, Quebec, Canada. Fic. 12. Foerstediscus grandis, new genus and species. The holotype, X 2, a flat circular expansion with the ambulacra all directed toward the right. Trenton (Curdsville limestone): Near Troy, Woodford County, Ky. L835 INO 334) fPLo 4] VOL. SMITHSONIAN MISCELLANEOUS COLLECTIONS = FP nse x % ¥ pedeergll si tif i hii Mh a 10 EDRIOASTEROIDEA Re LO) g see pa (For explanation, SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 9 NEW SPECIES OF TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA (Wirth NINE PLATES) BY FERDINAND CANU AND RAY S. BASSLER Head Curator, Department of Geology, U. S. National Museum Try ee, E-INC ZOE VIG, fi oo “ANo 1 = (PUBLICATION 3302) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION APRIL 26, 1935 The Lord Baltimore Press BALTIMORE, MD., U. 8. A. NEW SPECIES OF TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA By FERDINAND CANU AND RAY S. BASSLER Head Curator, Department of Geology, U. S. National Museum (With NINE PtrateEs) INTRODUCTION Through the kindness of various Australian friends, particularly Dr. F. Chapman, I have from time to time received many packages of washings containing Tertiary bryozoans from Victoria, with the re- sult that as our studies upon other faunas progressed, Dr. Canu and I were able not only to classify more accurately, and make new ob- servations upon, the known Victorian species but also to bring to light a number of interesting new forms. By the fall of 1931 such a volume of notes had accumulated that Dr. Canu began their compila- tion for publication. On February 11, 1932, he completed the editing and forwarded the final pages of these notes to me. The next morning, awakening with a slight headache, which grew steadily worse, he passed away from cerebral hemorrhage within a few hours. Thus came to a close our association in scientific work of almost a quarter of a century. Economic conditions have prevented the publication of our com- plete work upon this subject, and the following abridged descrip- tions of the new species are issued at the present time to make them available for stratigraphic use in South Australian geology. Our studies would seem to indicate that the Australian Tertiary does not cover a long time period. The bryozoan faunas are so unlike the standard associations in the Tertiary of Europe and America that no definite correlations have been made so far. Indeed, their nearest re- lations seem to be in the recent seas around Australia. All the types of the species here described are in the collections of the United States National Museum. RS: BASSEER. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 93, No. 9 bo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Class BRYOZOA Ehrenberg Order CHEILOSTOMATA Busk Suborder Anasca Levinsen Division MALACOSTEGA Levinsen, 1909 Family MEMBRANIPORIDAE Busk, 1854 ACANTHODESIA Canu and Bassler, 1920 ACANTHODESIA QUADRILATERA, n. sp. Plate 1, figs 10 Description—The zoarium is subcylindrical, consisting of six rows of zooecia arranged around a central line. The zooecia are distinct, adjacent to each other through their mural rims, elongated, rectangu- lar. The mural rim is thin, rounded, granulated, common to the adjacent zooecia; the cryptocyst is flat, smooth, shorter than the opesium. The opesium is anterior, elongated, elliptical, regular, bor- dered by a salient cushion, ornamented by small, very short spicules. Measurements.— ho=0.27 mm Lz=0.48 mm Opesium Zooeciu lo=0.12 mm lz=0.25 mm Occurrence-—Muddy Creek (Balcombian), and Orbost, Gippsland (Janjukian), Victoria. Holotype -—U.S.N.M. nos. 85582, 85583. ACANTHODESIA REGULARIS, n. sp. Plate 1, fig. 3 Description—Zoarium of slightly flattened, subcylindrical branches of 6 to 10 rows of zooecia arranged around a central line. Zooecia rectangular with granulated mural rim and a cryptocyst about three fifths as long as the opesium, which is anterior and elongated. Measurements.— ho=0.30 mm Lzg=0.60-0.65 mm Opesium{ lo=0.18-0.20 mm Zooeciay lz=0.30-0.35 mm Occurrence.—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes—U.S.N.M. no. 85581. NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER iS MEMBRANIPORA Blainville, 1830 MEMBRANIPORA AREOLATA, n. sp. Plate 1, fig. 2 Description—The zoarium is bilamellar; the fronds have four or five longitudinal rows of zooecia on each face. The zooecia are distinct, separated by a furrow or by a quadrangular area more or less broad, a little elongated, elliptical. The mural rim is thin, flat, of irregular width, smooth, and bears distally a small transverse triangular avicu- larium with two denticles and a pointed beak. The opesium is large, of the same form as the zooecium. The quadrangular area is very irregular in form and position. Ovicell unknown. Measurements.— Ore ho=0.40-0.50 mm z ; Lz=0.60 mm m1 ooecium Pesiun) Jo=o. 35 mm lz=0.45-0.50 mm Occurrence.—Janjukian ; Aire Coastal Beds, Victoria. Holotype —U.S.N.M. no. 60153. VINCULARIA Defrance, 1829 VINCULARIA GIGANTEA, n. sp. Plate 1, fig. 4 Description —The zoarium is rodlike (vincular) in form, composed of four or five longitudinal rows of cells arranged around a central line. The zooecia are distinct, gigantic, much elongated, rounded distally, narrowed proximally. The mural rim is thin, smooth, salient, rounded; the cryptocyst is very large, flat, smooth, or very finely granulated ; the opesium is elliptical, elongated, margined by a salient swelling, terminal, and much smaller than the cryptocyst. Measurements.— ho=0.50-0.65 mm ; L220) fim lo=0.30 mm BeUee lz=0.65 mm Occurrence—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes—U.S.N.M. no. 85905. Opesium OTIONELLA Canu and Bassler, 1917 OTIONELLA CIRCUMDATA, n. sp. Plate 1, figs. II, 12 Description—The zoarium is a small, truncated, solid cone; the base is slightly convex and ornamented with sinuous radial lines; at 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 the periphery there are small elliptical cavities equally spaced, 11 or 12 in number, each containing a small avicularium. The zooecia are distinct, separated by their mural rim, arranged in radial rows, hex- agonal. The mural rim is thin, salient; the cryptocyst is large, con- cave, deep, smooth; the opesium is terminal, orbicular, margined by a salient thread. The vibracula are very large, auriform, primoserial. The zooecia are not closed around the apex. Measurements.— ho=0.07 mm { L2=0.25-0.30 mm lo=0.07 mm | lz=0.25-0.30 mm Occurrence.—Balcombian beds at Muddy Creek, near Hamilton, Victoria. Holotype —U.S.N.M. no. 85807. Opesium Family HINCKSINIDAE Canu and Bassler, 1927 HINCKSINA Norman, 1903 HINCKSINA UNISERIALIS, n. sp. Plate 2, fig. 9 Description—The zoarium incrusts bryozoa. The zooecia are ar- ranged in uniserial, ramified branches ; they are large, much elongated, pyriform, and bear proximally along caudal gymnocyst. The opesium is large, anterior, oval; the mural rim is salient, beveled, a little en- larged at the base and ornamented with spines. The ovicell is endo- zooecial, rather large, convex, smooth. Measurements.— ho=0.40-0.45 mm Lz=0.55-0.80 mm lo =0.20-0.25 mm lz=0.35-0.40 mm Occurrence.—Janjukian beds at Corio Bay, Geelong, Victoria. Holotype —U.S.N.M. no. 85756. Opesium Zooecium Family ALDERINIDAE Canu and Bassler, 1927 MEMBRANIPORIDRA Canu and Bassler, 1927 MEMBRANIPORIDRA (?) ASYMMETRICA, n. sp. Plate 1, fig. 9 Description—The zoarium is bilamellar. The zooecia are large, distinct, separated by a very deep furrow, elongated, elliptical; the gymnocyst almost entirely surrounds the opesium; it is convex and salient on the lateral parts, concave, deep, and hidden proximally. The mural rim is a thin, sinuous thread bearing a salient, simple, or NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 5 bifurcated apophysis on one side only ; the opesium is elongated, ellip- tical, asymmetrical. The proximal gymnocyst bears the ovicell or a large, transverse, inconstant, elliptical avicularium. The ovicell is hyperstomial, globular, smooth, closed by the opercular valve. Measurements.— ho=0.37-0.45 mm : Lg=0.75 mm lo=0.15-0.20 mm ——_ lz =0.30-0.40 mm Occurrence-—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85794. Opesium| ELLISINIDRA Canu and Bassler, 1933 ELLISINIDRA PYRIFORMIS, n. sp. Plate 4, fig. 7 Description—The zoarium incrusts shell fragments. The zooecia are distinct, separated by a very shallow furrow, little elongated, elliptical, or almost transverse with a general pyriform aspect. The mural rim is very thin, filiform, rounded, smooth; it bears distally a small triangular, transverse avicularium; the opesium is very large and of the same form as the zooecium. The ovicell is small, convex, hyperstomial, closed by the operculum. Measurements.— ho=0.35-0.40 mm Te Lz=0.45-0.50 mm Jo=0.30-0.40 mm lz =0.40-0.50 mm Occurrence.—Janjukian beds at Flinders and Mount Gambier ; Balcombian beds at Muddy Creek, Victoria. Cotypes—U.S.N.M. nos. 85691-85693. Opesium STAMENOCELLA Canu and Bassler, 1917 STAMENOCELLA FUSIFORMIS, n. sp. Plate 1, fig. 8 Description—The zoarium is free, cylindrical. The zooecia are distinct, separated by a common mural rim, very elongated, fusiform; the gymnocyst is short, rectangular, very little convex, smooth. The mural rim is thin, rounded, smooth; the opesium is very much elon- gated, elliptical, terminal. —The gymnocyst bears sporadically an orbic- ular avicularium. The ovicell is unknown. Measurements. ho=0.45 mm : fz—070 mm esium Zooecium oe | Jo=0.15-0.18 mm lgz=0.25 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype-—U.S.N.M. no. 85881. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 ALLANTOPORA Lang, 1914 ALLANTOPORA CONFINIS, n. sp. Platetr, ie 5 Description—The zoarium incrusts shells and forms uniserial or pluriserial rows of zooecia. The zooecia are distinct, separated by a deep furrow, very elongated pyriform, simply adjacent in the multi- serial portion ; the gymnocyst is large, very convex, smooth, narrowed proximally, forming a caudal portion. The mural rim is thick, rounded, smooth. It bears exteriorly to the termen a row of large spines en- tirely surrounding the opesium.' Ovicell and ancestrula unknown. Measurements.— ho=0.25-0.30 mm. ces Lg=0.60-0.75 mm lo =0.18-0.20 mm [z=0.30-0.40 mm Number of zooecia in 4 mm* (=4 square millimeters), 20-22. Occurrence-—Balcombian beds at Muddy Creek and Janjukian at Flinders and Mount Gambier, Victoria. Cotypes—U.S.N.M. nos. 85598-85600. Opesium AMPHIBLESTRUM Gray, 1848 AMPHIBLESTRUM GRANDE, n. sp. Plate 1, fig. I Description—The zoarium is bilamellar. The zooecia are distinct, separated by a deep furrow, very large, little elongated, ogival, sur- rounded on three quarters by a granular gymnocyst. The mural rim is salient, very thin distally, enlarged laterally, much attenuated proxi- mally ; the cryptocyst is shallow, flat, smooth, merging imperceptibly with the proximal gymnocyst ; the opesium is large, terminal, trans- verse, pyriform, slightly trifoliated. The ovicell is hyperstomial, not closed by the opercular valve, globular, smooth, margined by an ecto- oecium, resting on the distal zooecium. Measurements.— ho=0.30-0.35 mm Tobesiitn Loe ee mm lo=0.35-0.40 mm | Izg=0.50-0.75 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85602. Opesium RAMPHONOTUS Norman, 1894 RAMPHONOTUS (?) LAMELLOSUS, n. sp. Plate 2, fig. 1 Description—tVhe zoarium is bilamellar. The zooecia are distinct, separated by a shallow furrow, very much elongated, pyriform. The NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER ii mural rim is thin, salient, sharp, smooth ; the cryptocyst is flat, smooth, little deep, as long or longer than the opesium ; it bears on its proximal portion a large transverse, triangular avicularium, with denticles, and in which the beak is very pointed and often slightly curved; the opesium is elongated, elliptical, narrowed distally by two small con- dyles symmetrically placed. The ovicell is hyperstomial, placed on the cryptocyst of the distal zooecium, globular, smooth, crowned by a zooecial avicularium. Measurements.— f ho=0.20-0.25 mm Lzg=0.60-0.65 mm Opesiuin} gor Zooecium | (ess .13-0.15 mm 2 =0.32-0.35 mm Number of zooecia in 4 mm’, 21. Occurrence-—Balcombian beds at Muddy Creek and Janjukian at Dartmoor, Victoria. Holotype —U.S.N.M. nos. 85850, 85853. Family HIANTOPORIDAE MacGillivray, 1895 TREMOPORA Ortmann, 1890 TREMOPORA ORBICULATA, n. sp. Plate 2a fies Description—The zoarium is unilamellar; the dorsal bears fenes- trae. The zooecia are distinct, separated by a deep furrow in which the connecting tubes are visible, orbicular, or slightly elongated. The mural rim is thin distally and very much enlarged at the base; it bears two large distal spines and laterally a large elliptical avicularium placed very high, opposite which is situated a small avicularium or a short bifurcated spine. The opesium is elliptical or orbicular, of the same form as the zooecium. Measurements.— ‘ ho=0.50 mm Lzg=0.60-0.70 mm Opesinm{ lo=0.40-0.50 mm Zovecium) lz =0.60-0.65 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype—U.S.N.M. no. 85894. TREMOPORA STAMINIS, n. sp. Plate 2, figs. 4, 5 Description.—The zoarium is unilamellar, free. On the dorsal the fenestrae are short, linear, narrow, and separate the six more or less broad connecting tubes. The surface is convex, smooth, ornamented with small salient radicular pores. The zooecia are distinct, separated 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 by a deep furrow at the angles of junction; the mural rim is thin distally, somewhat enlarged at the base; it bears two small distal spines, simple or bifurcated. The avicularium is large, triangular, without pivot, placed on the mural rim and in the immediate vicinity of the opercular valve. The beak is salient outside of the cell, very slightly curved. At its base a large spine with numerous reticulated filamentous branches covers the zooecium and unites itself to the op- posite side of the mural rim; the lacunae are little numerous, long and linear. Always on the mural rim in front of the large avicularium there is a small, branching palmate spine placed above the last ramifica- tions of the large spine. The ovicell is hyperstomial, small, convex, granular. Measurements.— ‘ ho=0.60-0.70 mm_, _ (Le=075 mam Opesium| lo=0.25-0.30 mm ooecia| Iz=0.45-0.50 mm Number of zooecia in 4 mm’, 15. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype.—U.S.N.M. no. 85896. Family ARACHNOPUSIIDAE Jullien, 1888 ARACHNOPUSIA Jullien, 1886 ARACHNOPUSIA LINEARIS, n. sp. Plate 2, fig. 2 Description.—The zoarium is free and bilamellar. The zooecia are distinct, separated by a shallow furrow, very elongated, oval or el- liptical; the frontal is a pericyst perforated by large lacunae; the latter are irregular, orbicular or crescentric, arranged irregularly in transverse rows. The opesium is semielliptical, transverse ; the distal peristome is thin; the proximal lip is Jinear, thick, with a large inden- tation on one side; this indentation is the insertion of a large spine which has disappeared in fossilization. On the side opposite the in- dentation and adjacent to the peristome there is a small triangular avicularium, the beak oriented distally above each opesium, and ad- jacent to the peristome there is another small triangular avicularium, the beak of which is oriented proximally. Measurements.— ho=0.10 mm s he 6 ol eais Zooecium Number of zooecia in 4 mm’, 9-10. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85603. Lz=0.90-1.15 mm Oncnae Z=0.50 mm NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 9 Division COILOSTEGA Levinsen, 1909 Family OPESIULIDAE Jullien, 1888 RECTONYCHOCELLA Canu and Bassler, 1912 RECTONYCHOCELLA (?) DIMORPHOCELLA, n. sp. Plate 2, fig. 6 Description—The zoarium is bilamellar. The ordinary zooecia are distinct, separated by a furrow of little depth, elongated, ogival. The mural rim is little distinct and is confused with the cryptocyst; the latter is smaller than the opesium, concave in its vicinity, somewhat convex proximally, slightly granular; the opesium is terminal, large, pyriform, narrowed toward the top by two small lateral condyles symmetrically placed. The accessory zooecia are membraniporoid ; the elliptical opesium is surrounded by a rounded salient mural rim. Measurements.— : ho=0.35-0.40 mm Lz=0.65-0.80 mm Opesiam lo=0.30 mm Zooeciusn| lz =0.55-0.60 mm Number of zooecia in 4 mm’, 12. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85854. FLORIDINELLA Canu and Bassler, 1917 FLORIDINELLA AUSTRALIENSIS, n. sp. Plate 1, fig. 6 Description—The zoarium is free, bilamellar ; the fronds are nar- row and bear on each face four longitudinal rows of cells. The zooecia are distinct, separated by their common mural rim, elongated, ogival, much narrowed proximally. The mural rim is thin, salient, rounded ; the cryptocyst is large, shallow, concave, smooth; the opesium is terminal, elongated, semielliptical, with the proximal border straight or concave ; two small distal condyles, symmetrically arranged, slightly contract the opesium. Measurements.— ? ho=0.25-0.28 mm ; Lz=0.60-0.70 mm Opesium Zooecium lo=0.17-0.19 mm lz=0.40-0.43 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85722. IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 VIBRACELLA Waters, 1891 VIBRACELLA PARVULA, n. sp. Plate 3, figs. 8, 9 Description—The zoarium is a small truncated, solid cone; the base is ornamented with numerous radial ribs, dichotomously dividing, each containing one or two rows of large lunularian pores. The zooecia are distinct, separated by their mural rim, small, hexagonal, arranged in radial rows. The mural rim is thin, salient; the crypto- cyst is concave, shallow, smooth; the opesium is terminal, semielliptic, little elongated, the proximal border being a little convex with two lateral, shallow, opesiular indentations. The vibracula are large, auri- form, primoserial. Measurements.— Gast ho=0.10-0.1I2 mm 7 : L2=025 tm esium ooecium P lo=0.10 mm lz=0.25 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype.-—U.S.N.M. no. 85904. SELENARIA Busk, 1854 SELENARIA TRIFOLIATA, n. sp. Plate 2, figs. 7, 8 Description—TVhe zoarium is orbicular, cupuliform, thick, some- what convex ; the inner face is slightly concave, granular, costulated, with a row of large scattered pores in the middle of the radial ribs. The zooecia are distinct, adjacent through their mural rim, hexagonal, somewhat elongated or transverse. The mural rim is thin, salient; the cryptocyst is shallow, smooth, slightly concave. The opesium is slightly elongated, trifoliate; the distal portion is elliptical and bor- dered laterally by two triangular, very salient apophyses ; the opesiular portion is linear, transverse, with two deep and rounded lateral in- dentations. The vibracula are primoserial, very large, auriform, sepa- rated into two parts by a salient point. The ancestrula is large and surrounded by 10 much smaller zooecia, of which 5 are vibracula. Measurements.— : ho=0.27-0.30 mm Zooecia f{Lg=0.40-0.50 mm EE lo=0.24 mm (marginal ! = ginal) } /z=0.50 mm Occurrence.—balcombian beds at Muddy Creek, Victoria. Cotypes—U.S.N.M. no. 85863. NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER rE SELENARIA GRANDICELLA, n. sp. Plate 3, figs. 1, 2 Description—tThe zoarium is orbicular, cupuliform, little convex. The zooecia are distinct, separated by a shallow furrow, very large, hexagonal, elongated or transverse ; the cryptocyst is small, smooth, somewhat concave. The opesium is very large, trifoliate; the distal border bears a kind of vestibular arch; the lateral borders are some- times a little salient; the proximal border is convex; the opesiular indentations are wide, deep, rounded. The vibracula are very large, auriform, primoserial ; the inner distal portion is shallow. On the in- ner face of the zoarium there are broad radial ribs, convex, each perforated by two rows of large irregular lunularian pores. Measurements.— ho=0.35 mm Lz=0.50 mm lo=0.30 mm lz=0.40-0.50 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85864. Opesium) Zooecium Family STEGANOPORELLIDAE Hincks, 1884 SIPHONOPORELLA Hincks, 1880 SIPHONOPORELLA LIVINGSTONE], n. sp. Plate 3) fics 5 Description—Vhe zoarium is unilamellar. The zooecia are distinct, separated by a shallow furrow, elongated, elliptical, narrowed at the base; the mural rim is somewhat salient, slightly thickened, granu- - lated ; the cryptocyst is shallow, flat, granulated. The opesium is large, terminal, irregular ; the polypidian tube is very salient, oblique, wide. Measurements. — Diameter of polypidian tube, 0.12-0.15 mm Width of opesium, 0.35 mm : Lzg=0.75-1.00 mm Zooecinin ls=0.48 mm Number of zooecia in 4 mm’, 12. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85884. SIPHONOPORELLA FILIPARIETIS, n. sp. Plate 3, fig. 4 Description —The zoarium is unilamellar. The zooecia are distinct, separated by a deep furrow, elongated, ogival ; the mural rim is round, 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 very thin, filiform, salient; the cryptocyst is shallow, flat, very finely granulated. The opesium is large, terminal, semielliptical, transverse ; the polypidian tube is long, oblique, smooth, very narrow, expanded distally. Measurements.— Diameter of polypidian tube, 0.12 mm Width of opesium, 0.40-0.45 mm Lz=0.85 mm lz=0.45-0.50 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85883. Zooecium Family THALAMOPORELLIDAE Levinsen, 1902 THALAMOPORELLA Hincks, 1887 THALAMOPORELLA ELONGATA, n. sp. Plate 2, fig. 10 Description.—The zoarium is bilamellar; the fronds are narrow. The zooecia are distinct, separated by their mural rim, much elongated, narrow, subrectangular ; the mural rim is thin, rounded, salient, joined to the peristome ; the cryptocyst is concave, rather deep, finely granu- lated. The aperture is orbicular or a little transverse ; the peristome is thin, salient. The opesiules are wide, short, placed in the neighbor- hood of the aperture. Measurements — ha=0.12-0.15 mm la=0.15 mm Number of zooecia in 4 mm’, 21. Occurrence.—Janjukian beds at Mitchell River, Victoria. Holotype —U.S.N.M. no. 85897. Lz=0.70-0.75 mm Apertura Zooeciuin | Iz =0.25-0.30 mm Family ASPIDOSTOMATIDAE Jullien, 1888 MACROPORA MacGillivray, 1895 MACROPORA CLARKEI ATTENUATA, n. var. Plate 4, fig. 1 Description —The zooecia are separated by a furrow and not by a salient thread. Our specimens bear kenozooecia analogous to those figured by Waters, 1885. One of them bears traces of a broken ovicell. Occurrence—Janjukian beds at Flinders, Victoria. Cotype.—U.S.N.M. no. 85781. NO. Q - TERTIARY BRYOZOA—-CANU AND BASSLER 13 MACROPORA QUADRISERIATA, n. sp. Plate 3, fig. 3 Description—The zoarium is free, vincular, quadriserial. The zooecia are distinct, separated by a thin salient thread, elongated, hexagonal; the cryptocyst is large, little convex, perforated by nu- merous pores and ornamented with small tuberosities. The aperture is subterminal, large, semielliptical, transverse; the proximal border is straight with two small, lateral indentations ; the peristome is very thick and salient. At the base of each zooecium there are two polygonal areas outlined by the ramifications of a separating thread. Measurements.— ha=0.15-0.20 mm L2= 125 mm la=0.20-0.25 mm Zooecium 2=O75 smn Occurrence—Janjukian beds at Bairnsdale, Victoria. Holotype —U.S.N.M. no. 85705. Aperture} Division PSEUDOSTEGA Levinsen, 1909 Family CELLARIIDAE Hincks, 1880 CELLARIA Lamouroux, 1812 CELLARIA ORBICULARIA, n. sp. Plate 4, fig. 3 Description—The segments are quadrangular. The zooecia are distinct, separated by their mural rim, much elongated, hexagonal ; the mural rim is thick, rounded, regular, the cryptocyst is shallow, very little convex, finely granulated. The opesium is large, orbicular, without any proximal denticle, margined by a thick pad. The ovicell is endotoichal; it is closed by an orbicular lamella forming a large area above the opesium. Measurements.— {ha=0.20 mm Lz=0.90-1.00 mm ope aue la=0.20 mm — lz=0.45 mm Occurrence.—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Holotype —U.S.N.M. no. 85623. CELLARIA ATTENUATA, n. sp. Plate 4, fig. 4 Description—The segments are long and rounded distally. The zooecia are distinct, separated by a very thin salient thread, somewhat 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 elongated, hexagonal; the cryptocyst is convex and forms a shallow cavity in front of the aperture. The aperture is semielliptical, trans- verse; the proximal border is straight or a little concave, with two very small lateral indentations. The ovicell is endotoichal ; its orifice is a thin, crescentic slit. The avicularian zooecia have a large semi- elliptical, transverse opesium. Measurements. — ha=0.12-0.15 mm Lz=0.75-0.80 mm Aperture Zooecium ‘ P la=0.15-0.17 mm Iz=0.50 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85617. Family MEMBRANICELLARIIDAE Levinsen, 1909 OMOIOSIA Canu and Bassler, 1927 OMOIOSIA ELONGATA, n. sp. Plate sss nen Description—The zoarium is free, bilamellar. The zooecia are distinct, separated by their mural rim, elongated, hexagonal, arranged in alternating transverse rows ; the mural rim is thick, salient, triangu- lar in section; the cryptocyst is deep, concave, very finely granular and entirely surrounds the opesium. The opesium is large, elliptical, elongated, not adjacent to the mural rim, surrounded by a salient thread. The special zooecia are larger but of the same form; their opesium is elliptical, median, and measures 0.30 by 0.15 mm. Measurements — ho=0.20-0.22 mm lo=0.15 mm Number of zooecia in 4 mm’, 24. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype -—U.S.N.M. no. 86955. Lz=0.00-0.65 mm Opesium) lz=0.35-0.40 mm Zovecium) Division CELLULARINA Smitt, 1867 Family SCRUPOCELLARIIDAE Levinsen, 1g09 CRASPEDOZOUM MacGillivray, 1886 CRASPEDOZOUM (?) ELONGATUM, n. sp. Plate 3, figs. 6, 7 Description—The zoarium is free, unilamellar, the fronds being formed of three longitudinal rows of zooecia. The zooecia are dis- NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 15 tinct, separated by a common mural rim, very long, ogival, a little narrowed proximally. The mural rim is thick, salient, rounded; the cryptocyst is very large, deep, concave, smooth; the opesium is ter- minal, elongated, elliptical, very finely crenulated. The zooecia of the median rows alone bear a large avicularian chamber ; it is placed on the proximal cryptocyst where it occupies half of the length; it is rectangular and convex; the avicularium is median, small, salient, triangular, with the beak oriented proximally. The noncellular face of the fronds shows the limits of the lateral zooecia and their out- lines without any apparent relation to the series of median zooecia. Measurements. — ho=0.35-0.40 mm aes eee mm lo=0.25 mm lz=0.45 mm Occurrence-—Balcombian beds at Muddy Creek, Victoria. Cotypes—U.S.N.M. no. 85721. Opesium Suborder AscopHora Levinsen, 1909 Family CRIBRILINIDAE Hincks, 1880 CRIBRILINA Gray, 1848 CRIBRILINA CRASSICOLLIS, n. sp. Plate 1, fig. 7 Description—tThe zoarium is free, cylindrical, bifurcated, formed of four or five longitudinal rows of zooecia. The zooecia are distinct, separated by a furrow (when young), elongated, elliptical. A thick secondary calcification fills up the separating furrow joining together adjacent mural rims and surrounding the zooecium and the aperture. The frontal is convex, perforated by large lacunae arranged in quin- cunx. The apertural bar was rapidly covered with secondary calcifica- tion to form a thick peristome. The aperture is terminal, semielliptical, transverse, with a concave proximal border. Two small avicularia are arranged symmetrically on each side of the aperture. Measurements — ha=0.10 mm 2 le 0.05) min la=0.18 mm “O0°Cl4 Iz=0.45 mm Occurrence —Balcombian beds at Muddy Creek, and Janjukian beds at Batesford, Victoria. Cotypes—U.S.N.M. nos. 85664, 85665. 2 Aperture} 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 CRIBRILINA TERMINATA CORONATA, n. var. Plate 3, fig. 12 Description—tThe distal portion of the peristome is crowned with three or four avicularia. Measurements. — nes ha=0.15-0.20 mm 7 . {Lz=0.75-0.90 mm ertura ooecia P la=0.20 mm c lz=0.45-0.50 mm Number of zooecia in 4 mm’, 10-12. Occurrence—Balcombian at Muddy Creek, and Janjukian at Bairnsdale and Flinders, Victoria. Cotypes—U.S.N.M. nos. 85670, 85671. CRIBRILINA TRISERIATA, n. sp. Plate 3, fig. 10 Description—The zoarium is free, unilamellar and formed of only three rows of cells. The zooecia are distinct, separated by a furrow, very elongated, elliptical ; the mural rim is very thick, smooth, entirely surrounding the cell; the frontal is perforated by small lacunae sepa- rated by small granules. The aperture is semielliptical, transverse ; it is surrounded by a very thick peristome joined with the mural rim. Two oblique, triangular avicularia are arranged symmetrically on each side of the proximal border of the aperture. The lateral zooecia are oblique and longer than the axial zooecia. Measurements.— ha=0.11 mm ; Lz=0.90 mm RSLS la=0.16 mm Zocectam lz=0.45 mm Occurrence.—Janjukian beds at Flinders and Batesford, Victoria. Holotype —U.S.N.M. nos. 85668, 85660. Family PORINIDAE D’Orbigny, 1852 PORINA D’Orbigny. 1852 PORINA FISSURIFERA, n. sp. Plate 4, fig. 11 Porina gracilis (pars) MacGillivray, Trans. Roy. Soc. Victoria, vol. 4, p. 103, Dial zie TeOS. Measurements.— hp=0.14 mm : P Zooecium Ip=0.14 mm Number of zooecia in 4 mm’, 10-TT. Be=i.5/mm Peristomice lz=0.60 (?) mm NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER iy) Affinities —This species differs from Porina (Acropora) gracilis Milne Edwards, 1836, in its larger micrometric dimensions, in its ascopore, which is a longitudinal slit, 0.10 mm in length, decorated with two lateral lips, and in the presence of small spathulated latero- frontal avicularia. Occurrence —Balcombian beds at Muddy Creek, and Janjukian at Mount Gambier, Victoria. Holotype-—U.S.N.M. nos. 85584, 85585. PACHYTHECELLA Bassler, 1934 PACHYTHECELLA UNIFASCIATA, n. sp. Plate 4, fig. 5 Description—The zoarium is free, bifurcated; the branches are somewhat compressed and formed of two longitudinal rows of cells opening only on one side. The dorsal is convex and smooth, except that the outlines of the zooecia are marked off by ridges. The zooecia are indistinct, little elongated, smooth, the frontal is perforated by a large orbicular ascopore. The peristomie is little salient, rather long ; the peristomice is orbicular ; the peristome is thin, nonsalient. Measurements.— hp=0.15 mm ; Lz=0.45 mm Ip=0.15 mm 29seelta 2=0.30 (?) mm Occurrence—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85823. Peristomice PACHYTHECELLA ARMATA, n. sp. Plate 4, fig. 8 Description——The zoarium is free, bilamellar, formed of rather wide, compressed branches. The zooecia are indistinct, elongated, with very thick walls; the frontal is convex and covered with a large number of shallow pores. The peristomie is formed by the much thickened zooecial walls; the apertura is buried at the bottom of the peristomie and appears orbicular ; the peristomice is orbicular and little distinct because of the absence of the peristome. On the marginal zooecia the ascopore is replaced by a large triangular, nonsalient, trans- verse, oblique avicularium; its beak is oriented exteriorly. Measurements. — : . fhp=o.15 mm ; Peristomice Zooecium Ip=0.15 mm Number of zooecia in 4 mm’, 18. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85822. Lz=0.60-0.75 mm lI2=035 (7?) mim 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Family TUBUCELLARIIDAE Busk, 1884 TUBITRABECULARIA Canu and Bassler, 1934 TUBITRABECULARIA PRODITOR, n. sp. Plate 4, fig. 10 Description—The zoarium is articulated; the segments are large, clavate, somewhat compressed at their extremity. The zooecia are in- distinct, much elongated ; the frontal is convex, smooth, bordered by large pores, and formed by an epicalcification hiding the peristomie and supported by thin trabeculae radiating from the ascopore. The peristomie (when visible) is long, oblique, much reduced, smooth, tubular ; the peristomice is orbicular; the peristome is thin, smooth, sharp. The ascopore is small, tubular (when visible), placed at the base of the peristomie. The ovicelled zooecia are globular, salient, often grouped in variable numbers. Measurements.— Poagpemirey ns if - - te Number of zooecia in 1 mm’, 5. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85886. Lz=0.70-0.80 mm Zooecium lz=0.40 (?) mm Family PETRALIIDAE Levinsen, 1909 PETRALIELLA MacGillivray, 1887 PETRALIELLA VULTUR AVICULIFERA, n. var. Plate! 5, fig. 2 Description.—The zoarium is unilamellar. The zooecia are distinct, separated by a furrow, large, elongated, capitate; the frontal is con- vex and punctured by scattered pores. An avicularian umbo, very salient, very long and somewhat oblique, is placed before the aperture ; the mandible is very long and placed laterally. The aperture is large, suborbicular, a little transverse ; a small expanded lyrule is placed on the proximal border in the vicinity of two very short cardelles; the peristome is very thin, smooth, ornamented with four spines. On each side of the aperture there is a small elliptical avicularium, with pivot, oriented proximally ; sporadically on the longer zooecia there are one or two small elliptical avicularia with pivot oriented distally. NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 19 Measurements.— ha=0.16 mm ; la=0.20 mm Zooecia} Number of zooecia in 4 mm’, 12. Occurrence—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85825. Lz=0.75-1.00 mm A perture lz=0.40 mm PETRALIELLA TRACTIFERA, n. sp. Plate 5, fig. 1 Description—The zoarium is unilamellar, perhaps orbicular. The zooecia are distinct, separated by a salient thread or by a thickened band, large, elliptical, elongated, swollen; the frontal is a tremocyst with large expanded pores and bearing an orbicular, salient, avicu- larian umbo. The shield is a narrow circular band surrounding the aperture ; it bears from four to six large hollow spines. The aperture is large, circular ; it bears proximally, a small lyrule and two trans- verse cardelles always placed at the same level as the distal border of the lyrule; the peristome is thin, smooth, salient, strengthened ex- teriorly by the shield. On the dorsal face of the zoarium, the zooecia are distinct, subhexagonal, separated by thickened salient bands form- ing an interzooecial epicalcification. Measurements.— ha=0.15-0.20 mm _ {Lz=0.75-0.80 mm Aperture; la=0.15-0.20 mm cnecia | lz=0.60 mm Number of zooecia in 4 mm’, Io. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype. —U.S.N.M. no. 85824. PETRALIELLA (?) DENTICULATA, n. sp. Plate 4, fig. 9 Description—The zoarium is bilamellar with wide fronds. The zooecia are distinct, separated by a salient thread, very elongated, irregular ; the frontal is somewhat convex, granulose, bordered with areolar pores. The peristomie is long, the peristomice is oblique, very irregular, with proximal border denticulated by two or three salient rounded mucrons. The aperture is buried at the bottom of the peristomie and bears a well-developed lyrule. Lg=0.75-1.00 mm lz=0.40-0.50 mm Number of zooecia in 4 mm’, 15. . Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85826. Measurements.—Zooecium ! 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Family GIGANTOPORIDAE Bassler, 1935 GIGANTOPORA Ridley, 1881 GIGANTOPORA HYSTRIKX, n. sp. Plate 4, fig. 2 Description—The zoarium is free, cylindrical. The zooecia are distinct, separated by a large prominent thread, somewhat elongated, hexagonal; the frontal is convex, pierced by numerous small tremo- pores separated from one another by protruding spinelike granules. The peristomie is long, salient, partly buried on the distal zooecium ; the peristomice is oblique, elliptical, transverse. The spiramen is large (0.17 mm wide) crescentric, somewhat tubular. The two avicu- laria are small, triangular, almost transverse. Measurements. ; ) | ip=—O-10 mm ; Lz=0.85 mm Pesistomice} Ip=o.15 mm Zooecium gos nn Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85728. GIGANTOPORA HEXAGONALIS, n. sp. Plate 4, fig. 6 Description —The zoarium is unilamellar. The zooecia are distinct, separated by a salient thread, a little elongated, hexagonal ; the frontal is transversely concave, formed by a tremocyst with small pores sepa- rated by regular tuberosities. The peristomice is subcircular; the peristome is smooth, thin, salient. The spiramen is large (0.14 mm wide), crescentic, bordered by a distinct thread. The two avicularia are large, triangular, oblique, with pivot. The beaks join on the median zooecial axis above the spiramen and form an angle with thickened lines which partially cover the peristomice. Measurements.— Peristomice} ie ee a Number of zooecia in 4 mm’, 12. Occurrence.—Balcombian beds at Muddy Creek, and Janjukian at Flinders, Victoria. Holotype —U.S.N.M. nos. 85725, 85726. Lz=0.90 mm Zooecium lz=0.55 mm GIGANTOPORA PERFORATA, n. sp. Plate 5, fig. 10 Description—The zoarium is free, bilamellar, formed of fronds of varying width. The zooecia are large, little distinct, elongated ; NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 21 the frontal is little convex, punctured by rather large tremopores. The peristomice is large, orbicular or elliptical and elongated ; the peristo- mie is very short; the peristome is rather thick, smooth, little salient. The spiramen is a simple perforation in the short peristomie. The ovicell is hyperstomial, opening largely in the peristome, very large, globular, covered with scattered tremopores. The two avicularia are small, triangular, transverse or oblique, pointed downward. Measurements. hp=0.22-0.25 mm Ip=0.18 mm Number of zooecia in 4 mm’, 8-10. Occurrence.—Balcombian beds at Muddy Creek and Janjukian beds at Flinders, Victoria. Cotypes —U.S.N.M. no. 85729. Lz=0.90-1.00 mm Peristomice lz=0.65-0.70 mm Zooecinin GIGANTOPORA ELONGATA, n. sp. Plate 5, fig. 5 Description—The zoarium is free, cylindrical. The zooecia are dis- tinct, separated by a furrow bordered by two very thick salient threads, large, very elongated, sinuous ; the frontal is a tremocyst, little convex and penetrated by rather large pores. The peristomie is short; the peristomice is orbicular or elliptical and trarisverse; the peristome is smooth, thick, little salient. The spiramen is large (0.25 mm wide) simple, not salient, marginated, somewhat concentric. The two peristomial avicularia are large, triangular, with pivot; their beak is pointed, oriented toward the peristomice. Measurements.— hp=0.20 mm Ip =0.22-0.25 mm Number of zooecia in 4 mm’, 6. Occurrence —Janjukian beds at Gellibrand, Victoria. Holotype—U.S.N.M. no. 85727. f3= 1-25-0230 mim Peristomice Iz =0.65-0.75 mm Zooecia} GIGANTOPORA MINUTIPOROSA, n. sp. Plate 5, fig. 9 Description—The zoarium is unilamellar, thick. The zooecia are distinct, separated by a thick salient thread, elongated, subhexagonal ; the frontal is somewhat convex, pierced with very small pores sepa- rated by small granulations. The peristomie is short, somewhat buried in the distal zooecium ; the peristomice is large, orbicular or elliptical, and transverse, little oblique ; the peristome is salient, thick granulated, 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 or crenulated. The spiramen is large (0.20 mm wide) distally tubular, granulated, oblique, oriented proximally. The two frontal avicu- laria are relatively small, triangular, with pivot; the beak is rounded and united to the thickened proximal portion of the peristome. Measurements.— hp=0.22 mm ; Lzg=1.08-1.17 mm lp =0.22-0.27 mm a lz =0.63-0.72 mm Number of zooecia in 4 mm’, 7. Occurrence.—Janjukian beds at Flinders and Batesford, Victoria. Holotype —U.S.N.M. nos. 85730, 85731. Peristomice GEPHYROPHORA Busk, 1884 GEPHYROPHORA BILAMELLARIA, n. sp. Plate 5, fig. 3 Description—The zoarium is free, bilamellar ; the fronds are broad and very thick. The zooecia are distinct, separated by a salient thread, large, rectangular, somewhat elongated; the frontal is little convex, almost flat, and formed by a granular tremocyst with numerous pores. The apertura is large, oval, with a broad, rounded proximal sinus ; the peristome is thin and very little salient. The ovicell is very large, globular, of the same nature as the frontal. On each side of the apertura there is a large triangular avicularium with pivot, with beak very pointed and oriented obliquely toward the apertural sinus. Measurements.— {ha=0.20 mm » tLe =0.7.0 mm acd Ineo arin as jp oeomnin Number of zooecia in 4 mm”, ITI. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85723. SPIROPORINA Stoliczka, 1864 SPIROPORINA TENUIS, n. sp. Plate 5, fig. 4 Description—The zoarium is free, cylindrical, bifurcated, very thin, formed of only four rows of cells. The zooecia are distinct, separated by a very small and very finely crenulated thread, much elongated, subcylindrical; the frontal is convex, perforated by very small tremopores arranged in linear rows and separated by scattered granulations. The peristomie is little apparent and of variable length. The peristomice is elliptical, transverse, oblique; the peristome is NO. Q TERTIARY BRYOZOA—CANU AND BASSLER 23 rather thick, fimbriated or crenulated. The spiramen is a small perfora- tion placed in the vicinity of the peristomice. Measurements.— hp=0.08 mm ; Lz=0.45-0.50 mm Ip=0.15 mm a lz=0.25 mm Occurrence.—Balcombian at Muddy Creek and Janjukian at Corio Bay, Geelong, etc., Victoria. Holotype—U.S.N.M. no. 85738. Peristomice Family SCHIZOPORELLIDAE Bassler, 1935 Subfamily ScHIzoporELLAE Canu and Bassler, 1917 BUFFONELLODES Strand, 1928 BUFFONELLODES BACULINA, n. sp. Plate 5, fig. 11 Description.—The zoarium is free, cylindrical, formed of five or six longitudinal rows of zooecia. The zooecia are distinct, separated by a furrow, somewhat elongated, lozenge shape, wide; the frontal is con- vex, smooth. The apertura is elongated, oval, terminated proximally by a wide rounded sinus; the peristome is wide, smooth, little salient. Measurements.— ha=0.10-0.12 mm oS —=@O100-0.75 Tm 2a la=0.08 mm Zeocetats lz=0.50 fh Occurrence—Balcombian beds at Muddy Creek, and Janjukian beds at Boggy Creek, Victoria. Holotype —U.S.N.M. nos. 85606, 85607. STEPHANOSELLA Canu and Bassler, 1917 STEPHANOSELLA SEXSPINOSA, n. sp. Plate 6, fig. 1 Description—tThe zoarium is incrusting. The zooecia are distinct, separated by a furrow, rather large, wide, ensiform; the frontal is convex, smooth. The aperture is suborbicular; the proximal border bears a wide rounded sinus; the peristome is very thin, little salient, ornamented with six very short spines. On each side of the aperture, distant from the peristome, there is a distinct, cylindrical, avicularian chamber, terminated by an orbicular, oblique orifice. The ovicell is hyperstomial, not closed by the operculum, very salient, globular, granulose, ornamented with a large, smooth triangular area. On cer- tain zooecia, the aperture is hidden by a large flat mucron; it bears 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 very frequently the base of a very large transverse avicularium with spathulate pivot in which the beak is supported on the peristome of an adjacent zooecium. Measurements.— ha=o0.12 mm _ (including Aperture I lone mm _ oral sinus) : L2z=0.65 mm Zooecium / 5 | ls=0.40-0.55 mm Number of zooecia in 4 mm’, 16. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85885. DAKARIA Jullien, 1903 DAKARIA CRASSOCIRCA, n. sp. Plate 6, fig. 7 Description —The zoarium is free, cylindrical, formed by four longi- tudinal rows of zooecia. The zooecia are distinct, separated by a furrow, at the bottom of which is a salient thread, elongated, elliptical ; the frontal is very convex and covered with large infundibuliform tremopores. The aperture is transverse, semielliptical; the proximal border bears a very wide rounded sinus; the peristome is complete, very thick ; between the inner proximal portion and the aperture, there is a small armature of little depth and characteristic of the genus. Ovicell unknown. Measurements.— ha=0.12 mm : Lz=0.84 mm Aperture la=0.15 mm Zoecinn [eosin Occurrence.—Janjukian beds at Bairnsdale, Victoria. Holotype —U.S.N.M. no. 85683. SCHIZOBRACHIELLA Canu and Bassler, 1920 SCHIZOBRACHIELLA HEXAGONALIS, n. sp. Plate 6, fig. 5 Description—The zoarium is free, unilamellar. The zooecia are distinct, separated by a salient thread, hexagonal, somewhat elongated ; the frontal is convex and formed of a tremocyst with small, numer- ous pores. The aperture, orbicular in aspect, is formed by a large semicircular anter and by a concave poster notched by a wide, shal- low sinus separated from the anter by two small lateral indentations. Laterally, at the height of the apertural sinus there is either a longi- tudinal slit or a small elongated avicularium. NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER ho on Measurements. — ha=0.15-0.17 mm la=0.17 mm Number of zooecia in 4 mm’, IT. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85861. Lz=0.80 mm Aperture lz=0.50-0.55 mm Zooecinn | CHIASTOSELLA Canu and Bassler, 1934 CHIASTOSELLA LAMELLATA, n. sp. Plate 6, fig. 8 Description—TVhe zoarium is free and lamellar; the fronds are more or less wide, bifurcated. The zooecia are little distinct, vaguely separated by a white line irregularly placed between the areolar pores, elongated ; the frontal is narrow, bordered by two scattered rows of areolar pores concealing the little apparent pleurocyst. The aperture is semicircular ; the proximal border is rectilinear and notched by a straight, short sinus rounded at its extremity; the peristome is thick, little salient, with four or five large distal spines. The ovicell is large, embedded in the distal zooecium, hyperstomial, not closed by the operculum; the ectooecium is large, circular, convex, smooth, orna- mented on the periphery with lines of small pores arranged radially. The two avicularia are arranged transversely on the transverse median axis of the zooecium ; they are long and thin with pivot, very pointed, projecting on the adjacent zooecia ; their base is placed on the exterior line of areolar pores. Measurements.— ha=0.15 mm la=0.10-0.12 mm Number of zooecia in 4 mm’, 16 to 20. Occurrence—Balcombian beds at Muddy Creek, and Janjukian at Mount Gambier, Victoria. Cotypes —U.S.N.M. nos. 85630, 85631. Aperture Zooecium | Lz=0.60-0.65 mm CHIASTOSELLA GIBBERA, n. sp. Plate 6, fig. 2 Description—The zoarium is unilamellar. The zooecia are little distinct, elongated, irregular ; the frontal is convex, ornamented with a longitudinal, median gibbosity surrounded by a double row of scattered areolar pores often separated by pseudocostules. The aper- ture is semielliptical, transverse; the proximal border is somewhat concave and bears a very narrow linear sinus; the peristome is non- 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 salient and bears four inconstant spines. The zooecial avicularium is transverse, thin with pivot, nonsalient, inconstant. The ovicell is unknown. Measurements. — x ha=oO.1 2) mm: f J Lz=0.60 mm DEORE la=0.15 mm Be Iz =0.25-0.30 mm 3 Number of zooecia in 4 mm’, 18-20. Occurrence.—Janjukian beds at Bairnsdale, Victoria. Cotype —U.S.N.M. no. 85629. CHIASTOSELLA POROSA, n. sp. Plate 6, fig. 4 Description—The zoarium is free, unilamellar. The zooecia are distinct, separated by a furrow, ogival, wide, often transverse; the frontal is convex, very porous, with much reduced pleurocyst. The aperture is suborbicular; the concave poster bears a rounded sinus of little depth; the peristome is wide, nonsalient, with three or four large spines. The avicularium is placed transversely on the median axis of the zooecium; it is very long, with pivot, with a beak thinned and placed on a convex, porous chamber. Ovicell unknown. Measurements.— ha=0.16-0.20 mm la=0.20-0.22 mm Number of zooecia in 4 mm’, 12. Occurrence—Balcombian beds at Muddy Creek, Victoria. Holotype—U.S.N.M. no. 85632. E2z=0.70 mm Aperture P lz=0.80 mm Zooecium i CHIASTOSELLA GRANDICELLA, n. sp. Plate 6, fig. 3 Description—The zoarium is free, unilamellar. The zooecia are distinct, separated by a furrow, very large, ogival, very wide, trans- verse ; the frontal is convex, porous, bordered by three rows of areolar pores leaving only a small frontal pleurocyst. The aperture is large, suborbicular ; the proximal border is very concave with a wide, rounded, rather deep sinus; the peristome is very thick, nonsalient and provided with four large distal spines. The zooecial avicularium is very large, rather long, with pivot, arranged transversely; the beak is pointed. Ovicell unknown. Measurements.— eee ha=0.22-0.25 mm 7 ’ Lz=0:90 mm erture ooecium ; P la=0.22 mm lz=0.80 mm (ir- regular ) Number of zooecia in 4 mm’, 5. NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 27 Occurrence-—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85628. CHIASTOSELLA PARVIPOROSA, n. sp. Plate 6, fig. Io Description——tThe zoarium is unilamellar. The zooecia are distinct, separated by a deep furrow, very large, ogival, little elongated, wide ; the frontal is convex, covered with relatively small pores, without pleurocyst. The aperture, transverse and semielliptical in aspect, bears on its concave proximal border a small rounded sinus; the peristome is very thick, nonsalient, garnished with four large hollow spines. Ovicell unknown. The avicularium is large, transverse, borne on a large porous convex chamber ; the beak is rather pointed; its base is placed on the first row of pores. Measurements.— ha=o0.28 mm f Lz=0.50-0.60 mm Aperture | la=0.25-0.27 mm Zoveciuen | Iz=0.50-0.55 mm (without avicularium) Number of zooecia in 4 mm’, 5. Occurrence —Janjukian beds at Flinders, Victoria. Holotype —U.S.N.M. no. 85627. EMBALLOTHECA Levinsen, 1909 EMBALLOTHECA INCLINATA, n. sp. Plate 6, fig. 6 Description—The zoarium is bilamellar, with broad fronds. The zooecia are distinct, separated by a white nonsalient thread ; the frontal is flat, perforated by large polygonal, expanded tremopores, rectangu- lar and somewhat elongated. The aperture is semielliptical, transverse ; the proximal border bears a semicylindrical mucron, inclined in the aperture; two small lateral cardelles are placed at the level of the distal border of the mucron ; the peristome is incomplete, wide, smooth, nonsalient. The ovicells are enormous, embedded in the distal zo- oecium, very convex and salient, covered with large pores. The ovi- celled zooecia are wider; their aperture is large (0.20 by 0.25 mm) and ornamented with two large cardelles; the peristomice is semi- elliptical, transverse. Measurements. — ha=0.10 mm Lz=0.60-0.70 mm la=0.15-0.16 mm 2NC SD Iz=0.40-0.50 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85701. Aperture} 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 EMBALLOTHECA GRANULATA, n. sp. Plate 5, fig. 7 Description——The zoarium is bilamellar. The zooecia are distinct, separated by a deep furrow, rectangular, elongated, a little contracted behind ; the frontal is convex, covered with granules separated by very small tremopores. The aperture is semielliptical, transverse ; the proxi- mal border is formed by a wide convex mucron; there are two car- delles, long, thin, oblique, oriented proximally ; the peristome is thin, smooth, hardly salient. The ovicell is unknown. Measurements. — ha=o.10 mm ] la=0.18-0.20 mm Zooecium) Number of zooecia in 4 mm’, 13, 14. Occurrence.—Kalimnan beds (bore no. 1, depth 110 feet) at Lakes I’ntrance, Victoria. Holotype —U.S.N.M. no. 85703. Lz=0.75-0.90 mm Aperture 2=0.40-0.50 mm EMBALLOTHECA ANGUSTATA, n. sp. Plate 5, fig. 8 Description—The zoarium is bilamellar. The zooecia are distinct, separated by a thin thread placed at the bottom of a furrow, much elongated, very narrow, of cylindrical aspect; the frontal is convex, finely granulose and perforated by a large number of very small tremopores. The aperture is semicircular; the proximal border is formed by a wide convex mucron presenting a small circular depres- sion on the zooecial axis ; the peristome is incomplete, smooth, salient. There are two thin cardelles. Measurements.— (ha=0.15 mm {eon mm Number of zooecia in 4 mm’, 13. Occurrence—Janjukian beds at Orbost, Gippsland, Victoria. Holotype —U.S.N.M. no. 85704. = 0.80-0.95 mm Aperture =0.35 mm Zooecium i ae L lg SCHIZOPORELLA Hincks, 1877 SCHIZOPORELLA ORBICULIFERA, n. sp. Plate 6, fig. 9 Description—The zoarium is free, cylindrical, formed of four or five longitudinal series of zooecia. The zooecia are distinct, separated by a very salient thread, lozenge-shaped, very much elongated, large ; NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 29 the frontal is convex, formed by a granular tremocyst with very small and numerous pores. The aperture is semielliptical, transverse; the proximal border bears a small rounded sinus; the peristome is com- plete, thin, smooth, little salient and separated from the separating thread only by a small furrow. The avicularium is small, orbicular, a little salient, always placed in one of the two lateral angles. The ovicell is large, globular, placed on the distal zooecium, hyperstomial, closed by the operculum; its surface is granular. The aperture of the ovicelled zooecia is larger. Measurements.— ha=0.10-0.12 mm : Lzg=0.90-1.00 mm AES lo=O0.15. mm Bouse Iz=0.50-0.60 mm Occurrence.—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes—U.S.N.M. no. 85856. SCHIZOPORELLA MACGILLIVRAYI, n. sp. Plate 9, fig. 5 Schizoporella phymatopora MacGillivray (pars), Trans. Roy. Soc. Victoria, vol. 4, p. 80, pl. 11, fig. 3 (not 2), 1895. Measurements.— oe ha=0.16 mm © 7 ; 2=0.77-0.85 mm rtur ium ee) Ja=O1620.07 mm yo Iz=0.44-0.55 mm Structure.—It is quite impossible that figures 3 and 2 of MacGil- livray refer to the same species, for there is a great difference in the apertural dimensions. We consider figure 3 as representing a distinct species, S. macgillivrayi, very close to S. alata, and characterized by its orbicular aperture (not transverse), its rounded proximal sinus, absence of avicularia, and zooecia frequently axially disarranged. Occurrence-—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85859. SCHIZOPORELLA TENUILAMELLOSA, n. sp. Plate 9, fig. 4 Description —The zoarium is bilamellar ; the fronds are flat, broad, and very thin. The zooecia are distinct, separated by a shallow fur- row, rectangular, very long; the frontal is a little convex, smooth, bordered with about 10 large areolar pores. The aperture in transverse aspect is semielliptical ; its proximal border is rectilinear and notched by a very small triangular sinus. Each zooecium bears on the median longitudinal axis of the frontal two very small avicularia; the first 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 is orbicular and placed a short distance from the apertural sinus ; the second placed a little lower, is transverse, very thin, triangular. The ovicell is unknown. Measurements.— fha=0.10 mm Piel co { | la=0.09-0.10 mm ™ Number of zooecia in 4 mm’, 15, 16. Occurrence —Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85860. Lz=0.65-0.75 mm Aperture lz=0.35-0.44 mm SCHIZOPORELLA PUSTULOSA, n. sp. Plate 5, fig. 6 Description—The zoarium is bilamellar with very narrow fronds. The zooecia are distinct, separated by a furrow, at the bottom of which is a very thin thread, elongated, somewhat oval; the frontal is a pleurocyst ornamented with seven to nine large granules or pustules and bordered by six very small areolar pores much separated from each other. The aperture is small, oval, a little oblique, em- bedded ; the proximal sinus is very wide, and triangular ; the peristome is very thin, nonsalient and bears four very small and much scattered tuberosities. Ovicell unknown. Measurements.— ha=o.10 mm . | Zooecium la=0.09 mm Number of zooecia in 4 mm’, 22-24. Occurrence.—Janjukian, Aire Coastal beds, Victoria. Holotype—U.S.N.M. no. 85862. Lzg=0.60-0.75 mm Aperture lz=0.35 mm SCHIZOPORELLA ARCANA, n. sp. Plate 9, fig. 9 Description—The zoarium is unilamellar. The frontal is not en- tirely perforated ; the cells are surrounded with an olocystic band and bear below the aperture a macula of the same nature. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Cotypes—U.S.N.M. no. 85857. SCHIZOPORELLA CLYPEATA, n. sp. Plate 6, fig. 11 Description—The zoarium is unilamellar. The zooecia are distinct, separated by a very thin thread, very large, nonsymmetrical, much NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 31 elongated, of little width; the frontal is little convex and formed by a large pleurocyst forming a smooth, elliptical cushion or shield; it is surrounded laterally by a double row of areolar pores and proxi- mally by four rows of scattered pores. The aperture is oval, axially disarranged, terminated by a narrow proximal sinus rounded at its extremity. A small oral avicularium, adjacent to the peristome is placed between the anter and the sinus. Another small zooecial avicu- larium is placed laterally on the exterior line of areolar pores and a little below the level of the proximal sinus. Ovicell unknown. Measurements. — ha=0.26 mm e240, mm Aperture Zooecia la=0.20 mm 2=0.65-0.75 mm Number of zooecia in 4 mm’, 6. Occurrence.—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Holotype —U.S.N.M. no. 85855. Subfamily ExocHELLINAeE Bassler, 1935 BATHOSELLA Canu and Bassler, 1917 BATHOSELLA LATICELLA, n. sp. Plate 9, fig. 2 Description —The zoarium is free, bilamellar, with narrow fronds. The zooecia are distinct, separated by a very deep furrow, elongated, globular, very wide; the frontal is rather convex, finely granular, pierced laterally by two or three pores. The aperture is large, ellipti- cal, transverse, somewhat oblique ; the peristome is thick, a little salient, smooth. On one zooecial margin there is a small triangular avicu- larium with pivot oriented distally. Measurements.— ha=0.14 mm la=0.20 mm Number of zooecia in 4 mm’, 12. Occurrence.—Janjukian, Aire Coastal beds, Victoria. Holotype —U.S.N.M. no. 60211. Lz=0.85 mm Zooecium [z=0.50-0.55 mm Aperture} BATHOSELLA BULBOSA, n. sp. Plate 9, fig. 1 Description—The zoarium is free, bilamellar, of narrow fronds. The zooecia are distinct, separated by a deep furrow, large, elongated bulbous ; the frontal is very convex, marginated by two to four large 3 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 pores, and formed by a granular pleurocyst. The aperture is large, elliptical, transverse, oblique, often mucronated ; the peristome is thick and granulated like the frontal. Laterally, on the transverse median axis of the zooecium and symmetrically arranged, there are two small elliptical, salient avicularia with pivot, oriented proximally. Measurements.— ha=0.12-0.15 mm : ' Zooecia la=0.20 mm Number of zooecia in 4 mm’, 14 or 15. Occurrence.—Janjukian, Aire Coastal beds, Victoria. Holotype-—U.S.N.M. no. 60205. Lz=0.75-0.80 mm Aperture lz=0.40-0.45 mm EXOCHELLA Jullien, 1888 EXOCHELLA GRANDIS, n. sp. Plate 9, fig. 3 Description— The zoarium is unilamellar. The zooecia are distinct, separated by a salient thread, large, elongated, hexagonal ; the frontal is a granular pleurocyst, bordered by large areolar pores separated by short costules. The apertura is semielliptical ; the peristome is thin and salient; a salient rectangular mucron hides the proximal border of the apertura. The ovicell is globular, convex, granular. The avicu- larium is transverse, thin, triangular, acuminated ; it replaces an areolar pore. Measurements.— ha=0.15-0.17 mm . (Lz=0.75-0.90 mm la=0.15-0.17 mm ooecla) ]z=0.50-0.70 mm Occurrence —Balcombian beds at Muddy Creek, Victoria. Holotype-—U.S.N.M. no. 85711. Apertura DIDYMOSELLA Canu and Bassler, 1917 DIDYMOSELLA CLYPEATA, n. sp. Plate 9, figs. 7, 8 Description—The zoarium is unilamellar, free ; the fronds are nar- row, bifurcated, formed of about seven longitudinal rows of cells; the dorsal is covered with an epicalcification hiding the form of the zooecia and on which there are deep longitudinal furrows. The zooecia are little distinct, separated by shallow and inconstant fur- rows, long aliform; the frontal is convex, formed distally by a shield, and proximally by a tremocyst with large pores. The shield is large, broad, smooth, perforated by two large foramina; its distal portion NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 33 is arched and separated by a deep furrow with two calcified bands at- tached to the peristome. The large zooecial avicularium is triangular, with pivot ; its beak is oriented exteriorly toward the nearest zooecial margin. The peristomice is semielliptical, transverse, with somewhat concave proximal border ; the peristome is thin, salient, ornamented with four spines, two of which are proximal. Measurements.— Peristomice| p _ ue oe) Number of zooecia in 4 mm’, 12. Occurrence—Muddy Creek (Balcombian), Anticline Creek, Dart- moor and Corio Bay, Geelong, Mount Gambier, etc., (Janjukian), Victoria. Holotype-—U.S.N.M. nos. 85684-85688. Lzg=0.72-0.75 mm Za ium Coe 2 =0.47-0.50 mm ESCHAROIDES Milne-Edwards, 1836 ESCHAROIDES ERECTA, n. sp. Plate 9, fig. 6 Description—The zoarium is free, erect, cylindrical, formed of four longitudinal series of zooecia. The zooecia are large, very elongated, somewhat aliform distally; the frontal is very convex, formed by a finely granulated pleurocyst, bordered by numerous areolar pores which separate very short costules; its distal portion is terminated by two large avicularian mucrons, very long, erect, hiding entirely the aperture and the locella. On each side of the aperture there is a small transverse, triangular avicularium in which the cham- ber is large, triangular, perforated in its middle by a small pore (radicell ?). Le= TA 5-25 mie lz=0.60 mm Occurrence.—Janjukian beds at Gellibrand, Torquay, Mount Gam- bier, and Boggy Creek, Victoria. Holotype —U.S.N.M. nos. 85819, 85847. Measurements —Zooecium Subfamily MicroporELLAE Canu and Bassler, 1917 MICROPORELLA Hincks, 1877 MICROPORELLA CAILLETI, n. sp. Plate 9, fig. 10 Description—The zoarium is unilamellar. The zooecia are distinct, separated by a deep furrow, hexagonal, wide, very little elongated ; the frontal is convex and is a finely granulated tremocyst perforated 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 by a large number of very small pores. The ascopore is transverse, elliptical or crescentric, salient and almost adjacent to the aperture. The aperture is semielliptical with a straight, transverse, proximal border ; the peristome is distal, salient, thick, ornamented with several flat, short spines. The avicularium is lateral, large, transverse, tri- angular, with beak pointed and oriented exteriorly, placed at the side or a little below the ascopore ; it surmounts a triangular chamber, salient, convex, covered by tremopores. The ovicell is hyperstomial, closed by the operculum, large, globular, of the same nature as the frontal. Measurements.— ha=0.07 mm : la=0.15 mm Zovecia} Number of zooecia in 4 mm’, 16. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85791. Le=0'55 mam Aperture 1z=0.40-0.55 mm FENESTRULINA Jullien, 1888 FENESTRULINA PRAETEXTA, n. sp. Plate 9, fig. 11 Description—The zoarium is free, bilamellar, bifurcated; the fronds are narrow and flabellate. The zooecia are distinct, bordered by a thin thread surmounting a large salient smooth cushion, triangu- lar in section, much elongated, rectangular somewhat narrowed be- hind; the frontal is little convex, bordered laterally by areolar pores and proximally with some tremopores; it bears on the longitudinal axis a large orbicular or crescentric ascopore much removed from the aperture. The aperture is semielliptical, transverse, the poster being simply concave. Ovicell unknown. Measurements. — ha=0.09 mm fete uibes saainal Number of zooecia in 4 mm’, 18. Occurrence.—Janjukian beds at Flinders, Victoria. Holotype —U.S.N.M. no. 85720. Lz=0.80 mm Zooecium a0 lz =0.30-0.40 mm Aperture! Subfamily HrppoporAr Canu and Bassler, 1917 HIPPOMENELLA Canu and Bassler, 1917 HIPPOMENELLA PARVIPOROSA, n. sp. Plate 7, fig. 2 Description—The zoarium is unilamellar and attached to frag- ments of bryozoa. The zooecia are distinct, separated by a rather deep NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 35 furrow, elongated, vaguely lozenge-shaped, enlarged on the transverse axis; the frontal is convex, smooth, bordered by a double row of numerous very small areolar pores separated by small radial costules. The aperture is oval, elongated ; the anter is separated by two cardelles descending from the poster, which is smaller. The frontal avicu- laria are small, triangular, with pivot; the beak is salient, pointed, oriented proximally ; they are placed indistinctly on the first or on the second row of pores; there are two or three on the same frontal. Measurements.— ha=0.20 mm ; so ae Zovecixm Number of zooecia in 4 mm’, 10. Occurrence—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85755. Lz=0.75-0.95 mm Aperture lz=0.60 mm HIPPOMENELLA RARIROSTRATA, n. sp. Blate a ties Description—tThe zoarium is unilamellar. The zooecia are distinct, separated by a deep furrow, elliptical, little elongated; the frontal is very convex, smooth, bordered by a double row of areolar pores. The aperture is large, semielliptical, elongated, with convex proximal border, without peristome. The ovicell is large, buried in the distal zooecium, hyperstomial, closed by the operculum; its frontal is deco- rated with numerous radial lines of small pores separated from each other by very salient costules ; the latter are arrested at the center by a large smooth, triangular area and a smooth, curved pad serves as the proximal limit. The frontal avicularia are rare, a single one, often absent, to a zooecium, small, triangular, placed very inferiorly on the inner line of areolar pores ; they have a pivot and the beak is oriented proximally. Measurements. — ha=0.18-0.20 mm ; Lz=0.66 mm peste la=0.16-0.18 mm 2 OUTS | lz=0.45 mm Number of zooecia in 4 mm’, 8. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype.—U.S.N.M. no. 85753. HIPPOMENELLA MAGNA, n. sp. Plate 8, fig. 13 Description —tThe zoarium is unilamellar. The zooecia are distinct, separated by a deep furrow, fusiform, large, very elongated; the frontal is convex and formed by a reduced pleurocyst, elliptical, elon- gated, bordered by two or three rows of lateral areolar pores and by 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 four or five rows of proximal ones. In the inner row the areolar pores are small, numerous, and separated by short radial costules. The aper- ture is neatly hippoporiform ; two long, oblique cardelles, placed low, separate a large orbicular anter from a poster smaller but wider and in which the proximal border is straight or a little convex ; no salient peristome. There is only a single lateral avicularium to a zooecium ; it is placed a little above the transverse, median zooecial axis ; it is small, oval, oblique, with beak oriented exteriorly. Measurements.— ha=0.22 mm la=0.29 mm Number of zooecia in 4 mm’, 6. Occurrence.—Janjukian beds at Torquay, 15 miles south of Geelong, Victoria (bore no. I at depth of 160 feet). Holotype —U.S.N.M. no. 85754. g=1.10-1.65 mm Aperture: Ig=0.55-0.65 mm Zooecium} HIPPOMENELLA VERMICULARIS, n. sp. Plate 8, fig. 3 Description —The zoarium is unilamellar, the zooecia are distinct, separated by a very deep furrow, elliptical, very short, swollen; the frontal is very convex and formed by a smooth pleurocyst, surrounded by four or five rows of large areolar pores. The aperture of the ovi- celled zooecia is rectangular, a little transverse, narrowed in the in- ferior third. The proximal border is convex. The ovicell is enormous, globular, embedded in the distal zooecium perforated by large scattered pores and richly decorated by salient vermiform ridges. There are two avicularia symmetrically arranged on the transverse median axis of each zooecium and on the inner line of areolar pores; they are small, triangular, oblique ; their beak is salient and is oriented exteriorly and proximally. Measurements — ha=0.15-0.20 mm , Lz=0.75 mm Aperture 5 Zooecium 75 la=0.21 tim lz=0.15 mm (Ovicelled zooecia) Number of zooecia in 1 mm’, 3. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85752. HIPPOPORELLA Canu, 1917 HIPPOPORELLA TESTU, n. sp. Plate 8, fig. 4 Description —tThe zoarium is unilamellar. The zooecia are distinct, separated by a deep furrow, large, elongated; the frontal is very NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 37 convex, smooth, bordered by a line of small areolar pores much scattered. The aperture is large, transverse, having the form of a dish cover with concave proximal border ; the peristome is incomplete, very little salient, thin, and bears six spines; the two proximal spines correspond to a slight lateral contraction of the aperture. The ovicell is large, globular, not closed by the operculum. A small oral avicu- larium placed on the line of the pores adjacent to the peristome is visible on one side of the aperture. Measurements.— ha=0.24 mm ; Ls= 1.00: mm la=0.30-0.33 mm a lz=0.60 mm Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85760. Aperture] HIPPOMONAVELLA Canu and Bassler, 1934 HIPPOMONAVELLA ACUTIROSTRIS, n. sp. Plate 7, fig. 5 Description—The zoarium is incrusting. The zooecia are distinct, separated by a salient thread, polygonal, elongated; the frontal is little convex, smooth, bordered by very small and numerous pores. The aperture is suborbicular, very little elongated ; the two cardelles are thin, salient, median ; the peristome is terminal, thin, little salient. The avicularium is thin, triangular, elongated, almost adjacent to the peristome ; the beak is very sharp and salient. Ovicell unknown. Measurements. — ha=0.14 mm (ee 2 mm Aperture lg=og mm asec scl ee ae ae Number of zooecia in 4 mm’, 13. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85757. Family SMITTINIDAE Levinsen, 1909 SMITTINA Norman, 1903 SMITTINA PERFORATA, n. sp. Plate 7tigs 3 Description—The zoarium is unilamellar. The zooecia are distinct, separated by a very small furrow, elongated, elliptical; the frontal is convex, marginated by a salient pad, convex and formed by a smooth pleurocyst surrounded by a line of six to eight pairs of large areolar pores separated by very short costules. The aperture is sub- 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 orbicular ; its proximal border bears a wide lyrule, salient and flat. In front of the aperture on the median longitudinal axis of the zo- oecium, there is a large orbicular perforation corresponding to an avicularium of the same form. Measurements. — ha=o0.14 mm la=0.12 mm Number of zooecia in 4 mm’, 13. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype -—U.S.N.M. no. 85868. Lz=0.63 mm Aperture Iz=0.41 mm Zooecium SMITTINA (?) PARVIOVICELLOSA, n. sp. Plate 7, fig. 4 Description—The zoarium is free, bilamellar, with wide, flat, or undulated fronds. The zooecia are distinct, separated by a salient thread, much elongated, fusiform; the frontal is convex, surrounded by small and very numerous areolar pores and formed by a rugose pleurocyst. The aperture is elliptical, transverse, without peristome. The ovicell is very small, globular, smooth, placed on the distal zo- oecium. The oral avicularium is small, adjacent to the aperture, el- liptical, with pivot, placed on the median longitudinal axis of the zooecium. Measurements. — ha=o.10 mm la = O21 mm Number of zooecia in 4 mm’, 12. Occurrence.—Janjukian beds at Orbost, Gippsland, Victoria. Holotype —U.S.N.M. no. 85869. Lzg=1.00-1.24 mm Aperture: lz =0.30-0.36 mm Zooecium | MUCRONELLA Hincks, 1880 MUCRONELLA ELONGATA, n. sp. Plate 7, fig. 6 Description.—The zoarium is free, cylindrical, rectilinear, of small diameter (0.80 mm). The zooecia are distinct, separated by a small, little salient thread, very long, narrow, fusiform. The frontal is convex, smooth, surrounded by a row of 14 to 16 pairs of very small areolar pores. The peristome is salient, thin, elliptical, elongated ; its proximal border bears a small mucron inclined in the peristomie ; the aperture is visible at the bottom of the peristomie. The ovicell is large, salient, globular, placed on the distal zooecium, smooth, hyper- stomial, opening in the peristomie. NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 39 Measurements — hp=0.15 mm Lg=1.15-1.25 mm Ip=0.12 mm [z=0.30-0.35 mm Occurrence.—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes.—U.S.N.M. no. 85806. Peristomice Zooecium SMITTINELLA Canu and Bassler, 1934 SMITTINELLA MAGNA, n. sp. Plate 8, fig. 7 Description—The zoarium is bilamellar with very narrow fronds. The zooecia are distinct, separated by a salient thread, large, rectangu- lar, much elongated ; the frontal is convex and perforated by three or four longitudinal rows of tremopores. The lateral zooecia are wider than the axial zooecia. The peristomice is suborbicular or elliptical ; the peristome is salient, thin, sharp, complete, with a small proximal spiramen. The ovicell is large, globular, marginated, finely granulated with a large porous area. Measurements. — hp=0.10-0.11 mm [p=0.10-0.11 mm } Lz=0.75-0.90 mm Zooecium Iz=0.25 mm (axial) 0.30 mm (lateral) Number of zooecia in I mm’, 3. Occurrence—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Holotype —U.S.N.M. no. 86957. Peristomice| SMITTINELLA OSIFERA, n. sp. Bilate7apicano Description—The zoarium is bilamellar with narrow fronds. The zooecia are distinct, separated by a small salient thread, somewhat fusiform, elongated ; the frontal is convex, finely granulose and per- forated by some scattered pores. The aperture is elliptical, transverse, with a very small proximal sinus ; the peristome is complete, very thin, scarcely salient. There is a broad lyrule and two small cardelles. The ovicell is very large, salient, globular with broad margin and a very fragile orbicular area; the orifice is very large, transverse, of the form of an open mouth, with a proximal spiramen. 4 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Measurements.— ha=0.12 mm. . {Lz=0.70-0.80 mm Aperture] la=0.17 mm cine | Iz=0.30 mm Number of zooecia in 4 mm’, 20-23. Occurrence.—Balcombian, beds at Muddy Creek, Victoria. Cotypes —U.S.N.M. no. 85871. SMITTINELLA MAGNIPOROSA, n. sp. Plate 7, fig. 8 Description—The zoarium is bilamellar with narrow, flat fronds. The zooecia are distinct, separated by a large salient thread, rectangu- lar, short, elongated ; the frontal is somewhat convex and perforated by a dozen large pores. The aperture is oval, a little elongated ; the proximal sinus is more or less wide, and always rounded; the lyrule and the cardelles are very fragile. The peristome is wide but not salient. The ovicell is large, globular with a central area orbicular and fragile. The peristomice of the ovicelled zooecia is large, semi- circular, transverse; the spiramen is large, salient, constant. Measurements.— ha=0.12-0.15 mm Aperture Zooecium © ig=0 82 mm Number of zooecia in 4 mm’, 25-30. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype—U.S.N.M. no. 85870. Lzg=0.50-0.55 mm lzg=0.22-0.25 mm PORELLA Gray, 1848 PORELLA TUBEROSA, n. sp. Plate 7, fig. 7 Description—The zoarium is bilamellar. The zooecia are distinct, separated by a salient thread, elongated, fusiform; the frontal is con- vex, covered by tremopores separated by tuberosities. The aperture is suborbicular; a hollow, rounded proximal indentation serves as orifice for an orbicular avicularium opening into the peristomice ; the chamber of this avicularium is large, smooth, and forms in front of the aperture a large gibbosity. The peristome is thin, little salient and forms a separating thread. The ovicell is very large ; a small axial ridge separates it into two compartments. The ovicelled zooecia are broader ; their aperture is enormous (0.15 by 0.25 mm). NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 41 Measurements.— ha=0.15 mm ‘ Lzg=0.75-1.00 mm Aperture] la=0.15-0.17 mm Zooeciunn} ‘= 0.29-0.34 mm Number of zooecia in 4 mm’, 15. Occurrence.—Balcombian, beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85843. PORELLA CYLINDROROSTRIS, n. sp. Plate 7, fig. 9 Description—The zoarium is bilamellar. The zooecia are distinct, separated by a thick thread, very salient, smooth; they are elongated, polygonal ; the frontal is little convex and perforated with large ex- panded tremopores. The peristome is thick, smooth, salient, covering proximally the avicularian chamber ; the peristomice is elliptical and transverse or suborbicular; into the peristomie opens a large cylin- drical avicularium, little salient, in which the orifice (invisible) is perpendicular to the apertural plane. The apertural chamber is large and forms a large convexity, salient, smooth, in front of the aperture. The ovicell is large, convex, marginated, with a fragile orbicular area ; it opens into the peristomie. Measurements — hp=0.13 mm lp=0.17 mm Number of zooecia in I mm’, 4. Occurrence.—Janjukian beds at Torquay near Geelong, Victoria. Holotype —U.S.N.M. no. 85829. Lz=0.70-0.77 mm Peristomice lg=0.35-0.42 mm Zooecium PORELLA BACULINA, n. sp. Plate 8 fig. 8 Description—The zoarium is free, cylindrical, in the form of a baton. The zooecia are distinct, separated by a small salient thread, elongated, elliptical, large; the frontal is convex and perforated by numerous small tremopores. The peristome is salient, thick, smooth, complete ; the peristomice is suborbicular; an elliptical avicularium is lodged in the proximal portion of the peristomie. Measurements. — hp=0.22-0.25 mm re es i2Z— TO) Tin Ip=0.25 mm lz=0.50-0.55 mm Occurrence.—Janjukian beds at Mitchell River, Bairnsdale, Mount Gambier, and Boggy Creek, Victoria. Cotypes—U.S.N.M. nos. 85830-85832, 85848. Peristomice| 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 PORELLA OPERCULATA, n. sp. Plate 8, fig. 6 Description—The zoarium is free, cylindrical, long. The zooecia are indistinct with thick walls, very long; the frontal is convex and covered with numerous tubular tremopores. The peristome is thick, smooth, very little salient; the peristomice is elliptical, elongated ; the peristomie is deep and formed by the thickening of the zooecial walls ; it contains in the proximal portion a large oblique avicularium, triangular, with pivot ; the beak is oriented distally ; the proximal por- tion is free or closed by a suborbicular, calcareous operculum. The ovicell is large, salient, globular, ornamented with a double row of large pores separated by short radial costules. Measurements.— hp =0.27-0.30 mm 4 L2=1.10-1.20 mm | lp=0.17-0.20 mm Bee iz=0.50 (7) mm Occurrence—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes—U.S.N.M. no. 85842. Peristomice PORELLA EXCAVATA, n. sp. Plate 7, fig. 11 Description— The zoarium is free, bilamellar ; the fronds are cylin- drical, flabellate or lamellar and narrow, bifurcated; there are from one to seven longitudinal rows of zooecia on each branch. The zooecia are distinct, separated by a salient thread, elongated, rectangular ; the frontal is flat and perforated with tremopores. The peristome is thin, salient, smooth; it is indented in the proximal portion by a linear sinus at the bottom of which there is a flat lyrule; the sinus is prolonged on the median axis of the zooecium by a small, linear excavation in which is placed a small, triangular, thin, elongated avicularium; the peristomice is orbicular and small. The ovicell is very small, globular, smooth. Measurements — hp=0.12 mm lp=0.12 mm Zooecium "jp oe 9 mm Iz=0.25-0.30 mm Occurrence—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes—U.S.N.M. no. 85838. Peristomice| NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 43 PORELLA RHOMBOIDALIS PARVIAPERTURA, n. var. Plate, 8, fig. 1 Description—Vhe aperture is small. There is a small round avicu- larium supported on a flat wide lyrule. The separating threads of the zooecia are very salient and thin. Measurements. — ; 1 f[hp=0.200 mm Pevistomice| Ip=0.18 mm Number of zooecia in 4 mm’, II. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85841. Lz=0.80-0.85 mm Zooecia 1 Iz=0.45 mm PORELLA RHOMBOIDALIS CRASSIMARGINATA, n. var. Plate 8, fig. 2 Description—tThe aperture is smaller than in the type. The small avicularium is borne on a flat lyrule. The separating threads of the zooecia are little salient and very thick. Measurements.— : . {hp=0.20 mm Peristomice} Ip=6.15 mm Number of zooecia in 4 mm’, Io. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Holotype —U.S.N.M. no. 85844. Lz=0.80 mm Zooecium sii lz=0.50-0.60 mm PALMICELLARIA Alder, 1864 PALMICELLARIA (?) MAGNA, n. sp. Plate. 8, ‘fie. 5 Description—tThe zoarium is free, very long, filiform, formed of four longitudinal rows of zooecia opposed two by two. The zooecia are distinct, separated by a small, little salient thread, very large, very long, tubulose; the frontal is very convex, fibrous and garnished laterally with numerous small areolar pores. The peristome is very long, oblique, rather salient, thin, with an inferior lip more developed and convex; the peristomice is large, semielliptical, transverse. An oral avicularium with basal chamber large and globular opens laterally on the interior of the peristomie. Measurements. — Peristomice} 7p 0.30 mm Lg=1.75-2.00 mm Zooecium Ip=0.30 mm lz=0.50 mm Qccurrence-—Janjukian beds at Anticline Creek, Dartmoor, Victoria. Cotypes. U.S.N.M. no. 85818. 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Family CELLEPORIDAE Busk, 1852 COSTAZIA Neviani, 1895 COSTAZIA CONVEXA, n. sp. Plate 8, fig. Io Haswellia producta (pars) MacGillivray, Trans. Roy. Soc. Victoria, vol. 4, p. 137, pl. 14, figs. 17, 18 (not 16 and 20), 1895. Description—The zoarium is cylindrical. The zooecia are distinct, poorly oriented, separated by a deep furrow; the frontal is very convex, almost tubular, smooth, surrounded by areolar pores, termi- nated by a mucron bearing an avicularium hiding a part of the aper- tura. The apertura is elliptical, somewhat elongated, without cardelles. The ovicell bears a perforated costulated area. The interzooecial avic- ularia are large, spathulated, enlarged at the beak, directed toward the base of the branches. Occurrence.—Balcombian. beds at Muddy Creek, Victoria. Cotypes—U.S.N.M. no. 85650. Family PHYLACTELLIDAE Canu and Bassler, 1917 PHYLACTELLA Hincks, 1880 PHYLACTELLA CHAPMANTI, n. sp. Plate 8, fig. 9 Description—The zoarium incrusts shells. The zooecia are dis- tinct, separated by a furrow, somewhat elongated, rather swollen ; the frontal is convex, smooth in appearance but perforated by extremely small pores and bordered by larger, more scattered pores. The aper- ture is small, semielliptical, with two cardelles and a lyrule visible only after suitably inclining the preparation ; the peristome is salient, somewhat thick. The ovicell is recumbent, small, globular, smooth, not closed by the operculum, opening in front of the oral mucron. Measurements. — ha=0.07 mm . {Lz=0.60-0.90 mm la=0.1I0 mm Zooecia =0.50-0.75 mm Occurrence.—Balcombian beds at Muddy Creek and Janjukian beds at Torquay (bore, 160 feet deep), Victoria. Holotype —U.S.N.M. nos. 85820, 85821. Aperture] NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 45 Family ORBITULIPORIDAE Canu and Bassler, 1923 STICHOPORINA Stoliczka, 1861 STICHOPORINA (?) PARVICAPITATA, n. sp. Plate 8, fig. 11 Description—The small fragments of this species in our collection are incomplete, and it is difficult to classify them. The ovicell which ought to be closed by the operculum, is small and analogous to that of Batoporaand Orbitulipora. We do not know if there was a central pit. The zooecial walls are olocystal with two or three pores at the base. On the interior face the zooecia are hexagonal, and without doubt this species is indeed one of the Orbituliporidae. Occurrence.—Balcombian beds at Muddy Creek, Victoria. Cotypes —U.S.N.M. no. 85882. EXPLANATION OF PLATES PLATE I (All illustrations on this plate are magnified, * 20) Fic. 1.. Amphiblestrum grande, new SPECiIES.........2.c00ceecccnscccsecs 6 Ovicelled fragment of a unilamellar zoarium illustrating the large size of the cells, the granular gymnocyst and the apparent ab- sence of avicularia. Balcombian beds at Muddy Creek, Victoria. Fic. 2. Membranipora areolata, new SpecieS........005.ccesccccccesscses 3 Portion of the bilamellar zoarium showing the distal transverse avicularia and the polygonal interopesial spaces. Janjukian, Aire Coastal beds, Victoria. Bug: “3. Acanthodesta regularis; new SPECIES... sc.ss us decease bee cde se 2 Portion of the free, subcylindrical branch exhibiting the regular form of the zooecia. Janjukian beds at Anticline Creek, Dartmoor, Victoria. BiG, A. Vaencularag: osgantéa; new iSPeCleS <.ccie esac whos « ceases 0s od cee 3 Portion of the rodlike zoarium composed of unusually large zooecia. Janjukian beds at Anticline Creek, Dartmoor, Victoria. Bias 5. 2dilantopora confints, new SpecieS <<. scuesscc sondeckc cscs cecae nc 6 A normal incrusting colony with zooecia bearing a large gymno- cyst. Balcombian beds at Muddy Creek, Victoria. Fic. 6. Floridinella australiensis, new species..........000.00ccccccuecee 9 The narrow bilamellar zoarium exhibiting the characteristic con- dyles in the opesium. Balcombian beds at Muddy Creek, Victoria. 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 PAGE Fie) 7 Gribrilina crassicollis, New Species: «eta eee iae 15 Surface of the free cylindrical zoarium. Balcombian beds at Muddy Creek, Victoria. Fie: 8; Stamenocella fustformis, new SpeCleS). «224. 2: e ee see eel cee 5 The free cylindrical zoarium celluliferous on all sides, exhibiting the fusiform zooecia and the orbicular avicularium. Balecombian beds at Muddy Creek, Victoria. Fic. 9. Membraniporidra asymmetrica, new SPECi€S...........00eeee evens Ordinary and ovicelled zooecia of the bilamellar zoarium. The spiniform process on one side of the zooecium and the large transverse avicularium are shown. Balcombian beds at Muddy Creek, Victoria. Fic. 10. Acanthodesia quadrilatera, new species..............0+-.eesseeee The delicate subcylindrical zoarium of elongated rectangular zooecia with short spicules indenting the opesium. Balcombian beds at Muddy Creek, Victoria. Eres: 1 02) (Ofonella cincumadata, NewasSpeClesa niece eieieieeiiee ieee tt. Lateral view of a small conical zoarium with large auriform primoserial vibracula. 12. Base of the same with radial lines and elliptical cavities each containing an avicularium around the periphery. Balcombian beds at Muddy Creek, Victoria. PLATE 2 (All illustrations on this plate are magnified, * 20) Fic. 1. Ramphonotus (?) lamellosus, new species................-..:-.. The bilamellar zoarium with normal and ovicelled zooecia and the narrow, transverse, triangular avicularium on the proximal part of the cryptocyst. Balcombian beds at Muddy Creek, Victoria. Bie. 2) Arachnopusia linearis. news Speclessadeeoaeeaee eee eee ce The free bilamellar zoarium showing the aperture with its recti- linear proximal border and the perforated zooecial frontal. Balcombian beds at Muddy Creek, Victoria. hic. 3. Lremopora orbiculata. New Species... ..a- cece eee eee etree Portion of the unilamellar zoarium illustrating the orbicular zooecia, the two distal spines and the large lateral avicularium with a small one opposite it. Balcombian beds at Muddy Creek, Victoria. Figs. 4, 5. Iremopora staminis, new SPECieS.....-. 6 .. sarees oe selec. 4. Frontal of a unilamellar zoarial fragment exhibiting ovicells and some zooecia with the large frontal spine preserved. 5. Dorsal of the same zoarium showing the fenestrae separating the connecting tubes, and the radicular pores. Balcombian beds at Muddy Creek, Victoria. Fic. 6. Rectonychocella dimorphocella, new species............0seeeeees Surface of the bilamellar colony with both ogival and membrani- poroid cells. 3alcombian beds at Muddy Creek, Victoria. iS) NI NI NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 47 HIGSIe OS CLEnGHIOUiTITOlalG, Mew; SPECIES waaa cite see ced os 6 cles see cis 10 7. Outer margin of an orbicular zoarium showing the labial eleva- tion of the opesium. 8. Inner side of a specimen with single row of large pores to each rib. Balcombian beds at Muddy Creek, Victoria. Bic. 0) Aincksmacunisertalis, new: SPeCieSs si. es. eee cle) sie os ve eels © 4 Part of the incrusting uniserial colony with one ovicelled zooecium. Janjukian beds at Corio Bay, Victoria. Fic. 10. Thalamoporella elongata, new species.................++20-000-- 12 The narrow bilamellar zoarium showing the much elongated zooecia and the inequality of the opesiules. Janjukian beds at Mitchell River, Victoria. PLATE 3 (All illustrations on this plate are magnified, < 20) PAGE Fics. 1, 2. Selenaria grandicella, new SPeCieS. 6... ..2< cn. cis sce ee owen II t. Cellular side of the cupuliform zoarium showing the vibracula with distal concavity. 2. Inner side of the same colony; the ribs have large lunularian pores. Balcombian beds at Muddy Creek, Victoria. Fic. 3. Macropora quadriseriata, new SPeCieS.......... 0.0 eeeseeeeeses 13 The quadriserial rod-shaped zoarium with zooecia ornamented by two polygonal areas at their base. Janjukian beds at Bairnsdale, Victoria. Fic. 4. Siphonoporella filiparietis, new SpecieS.......--...-..ee ee ee eens II Fragment of a unilamellar zoarium showing the thin mural rim, the narrow polypidian tube, and wide zooecia. Balcombian beds at Muddy Creek, Victoria. Fic. 5. Siphonoporella livingstonet, new SPeCieS............. eee ee renee II Unilamellar zoarium with narrow zooecia and wide polypidian tubes. Balcombian beds at Muddy Creek, Victoria. Fics. 6, 7. Craspedozoum elongatum, new SP€Ci€S..............eeeeeeee 14 6. Unilamellar, triserial fragment with the large avicularian chamber developed on the middle row. 7. Dorsal of the fragment shown in figure 6. Balcombian beds at Muddy Creek, Victoria. PIGS) Son. i soracciia. parvuld, NEW SPECIES aehscig sts < coi eee isieie ee alas 10 8. Cellular face of the small conical zoarium with large auriform primoserial vibracula and small hexagonal zooecia. 9. Base of the same with lunularian pores on the ribs. Balcombian beds at Muddy Creek, Victoria. Fic. 10: Gribrilina triseriata, New. SPECIES. «.. 2 ees. b ese eee cece eee ee cee 16 Frontal view of the unilamellar triserial zoarium. Janjukian beds at Flinders, Victoria. 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 PAGE Pic ri Omiotosta elongata, Mew SPECIES: acs masse ore nae nei eee ie oie eee 14 A bilamellar zoarium showing the elongate zooecia of two kinds— normal and special. Balcombian beds at Muddy Creek, Victoria. Fic. 12. Cribrilina terminata coronata, new variety........+-ssceeeeeseees 16 View of unilamellar zoarium illustrating small avicularia on the peristome. Balcombian beds at Muddy Creek, Victoria. PLATE 4 (All illustrations on this plate are magnified, 20) aie Fic. 1. Macropora clarkei attenuata, new variety..........00.eeeeeeee ees 12 Portion of the bilamellar zoarium showing the characteristic separating furrow. Janjukian beds at Flinders, Victoria. Gigantopora hystrix, new Species. 2... ssc. ee nee we cele enislere 20 Surface of the cylindrical zoarium showing the short, broad zooecia and salient spines separating the tremopores. Balcombian beds at Muddy Creek, Victoria. Picgoe3. Cellarianorbiculeria, Mew. SPECiESin.jesr ete seein ees 13 Portion of an ovicelled segment exhibiting the orbicular opesium. Janjukian beds at Anticline Creek, Dartmoor, Victoria. Ere) 4. Cellariaatienuata, mew Species’ as canst ieieee lente = adeteters yee de> 13 Extremity of a segment showing a joint of articulation at the end, a large broad avicularian zooecium, and the orifices of the ovicells. Balcombian beds at Muddy Creek, Victoria. Fie. 5. Pachythecella unifasciata, new Species ../\5. 2. j2ee accel teem ee 17 Celluliferous side of the unilamellar bifurcated zoarium showing the large orbicular ascopore. Balcombian beds at Muddy Creek, Victoria. Fic..6 Gigantopora hexagonalis, new SPeCieS........6--e aes secuencere cae 20 Several zooecia of the unilamellar zoarium illustrating the coales- cence of the beaks of the two large lateral avicularia above the spiramen. Balcombian beds at Muddy Creek, Victoria. Elltisimdra pyriformis;, NeW SPOCiES..f).). ste vile cere oiaie slefeterais es one 5 An example preserving the small hyperstomial ovicell. Balcombian beds at Muddy Creek, Victoria. Pic. 8: Pachythecella: armaia, new ‘SpeCieS. 3% 201. sais most esle ae sia ee 17 Bilamellar zoarium with orbicular aperture, small ascopore and the frontal of the zooecia covered with small shallow pores. 3alcombian beds at Muddy Creek, Victoria. Fic. 0. Petralielia (2?) denttculata, new: SpeCies. <0. ee 12 oe - sein elses ats 19 Surface of the bilamellar zoarium showing the denticulated border of the peristomice. Balcombian beds at Muddy Creek, Victoria. Fic. Ny Fic. N NO. 9 Fic. Io. RIG. tr RIG. i. EstGae 2: Fic. 3: Frc. 4: Fig, 5: LEE, (6) Fic. 7. Fic. 8. Fic. 9 TERTIARY BRYOZOA—CANU AND BASSLER Tubtirabecularta proditor, new SPeCieS..........2---es cece ceceee Portion of a complete ovicelled segment, showing trabeculae con- solidating the peristomies, the ascopore, and the tubular per- istomie and rounded peristomice. Balcombian beds at Muddy Creek, Victoria. IROTINOMASSUPUTE TUE W SPECIES larete rarer tele oicie aie syelel olaln/oin sauce a!» sherals Portion of a narrow bilamellar zoarium with large frontal avicu- laria and slitlike ascopore. Balcombian beds at Muddy Creek, Victoria. PLATE 5 (All illustrations on this plate are magnified, < 20) iRetaliella’ tractifera. TeWeSPCCIESay arc sies sioltelsleiee retain eile soe Zooecial surface of the unilamellar orbicular zoarium. Balcombian beds at Muddy Creek, Victoria. Petraliella vultur (Hincks, 1882) aviculifera, new variety........ Surface of the unilamellar zoarium showing the numerous small avicularia and the salient avicularian umbo with its laterally placed mandible. Balcombian beds at Muddy Creek, Victoria. Gephyrophora bilamellaria, new specieS...........----eeee ee eees Surface of the bilamellar zoarium. The zooecia are without spiramen. Balcombian beds at Muddy Creek, Victoria. SPIRO POMING sHENUES (IEW “SPECIES aco cisdye scsi so, dois at endne shs)e\ alas ees View of the thin cylindrical bifurcated zoarium composed of zooecia separated by a thin crenulated thread. Janjukian beds at Corio Bay, Victoria. Gigantopora elongaia: NeW SPECIES. 2/9). 5... 2 o.es eo 6 wien Pei ein waite bois The cylindrical zoarium exhibiting the elongate zooecia and the crescentric spiramen. Janjukian beds at Gellibrand River, Victoria. Scniwoporeiia: pustulosa, GEW SPECleS 5.20.2. s-e oe cesar enase os os Portion of the bilamellar zoarium showing the formation of the frontal pustules. Janjukian, Aire Coastal beds, Victoria. Emballotheca granulata, mew) Specicss..<22- sees ecesecs sc sss e soe Fragment of the bilamellar zoarium exhibiting the rectangular zooecia with frontal covered by granules. Kalimnan beds at Lakes Entrance, Victoria (bore no. 1 at depth of 110 feet). Emballotheca angustata, new SPECIES... 2.2.22 .06-.50c cess ceeces Fragment of the bilamellar zoarium with very narrow, much elongated zooecia. Janjukian beds at Orbost, Gippsland, Victoria. Gigantopora minutiporosa, MEW SPECIES............. cece wees ecco Celluliferous side of the unilamellar zoarium with the zooecial frontal pierced by minute pores. Janjukian beds at Flinders, Victoria. 16 22 22 2I 30 28 28 21 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Fig..10.. '(Gigantopora. perforata, Mew SPECIES... 2a 4.06 -. cave sie oe euninie ne 20 Surface of a narrow bilamellar ovicelled zoarium. Some zooecia have their peristomie complete and perforated by a small spiramen. Janjukian beds at Flinders, Victoria. Big. 1 Bunonellodes baculina, new, SPeCleS-- scene eee eee 22 Cylindrical unovicelled zoarium formed of zooecia with convex smooth frontal. Balcombian beds at Muddy Creek, Victoria. PLATE 6 (All illustrations on this plate are magnified, 20) PAGE Fic. 1. Stephanosella sexspinosa, new specieS................cceeeeeveee 23 Ovicelled incrusting zoarium with both zooecial and zoarial avicularia. Balcombian beds at Muddy Creek, Victoria. Fic. 20 Chiastosella gibbera: new Species. 2..-ce seca us on cee ree ae ee 25 Small fragment of a unilamellar zoarium illustrating the ir- regular indistinct zooecia and large falciform avicularia. Janjukian beds at Bairnsdale, Victoria. Fic. 3.. Chiastosella grandicella, new species..........2:+2+2.seeeneeeccs 26 Unilamellar zoarium exhibiting the very large zooecia and avicu- laria. Balcombian beds at Muddy Creek, Victoria. PiG.«14,° (Chiastosclla. porosa, Rew ISPECieS.20 wee een eae ee ee ee 26 Surface of the unilamellar zoarium showing the convex porous frontal and transverse avicularia. Balcombian beds at Muddy Creek, Victoria. Fic. 5. Schizobrachiella hexagonalis, new species...............eeeeeeeee 24 Zooecia of the unilamellar zoarium bearing only a single well- developed avicularium. Balcombian beds at Muddy Creek, Victoria. Bic. 16) “Emballotheca anclimata, mew sSpecies.n ee eee eee eee 27 Surface of the broad bilamellar type specimen showing the enor- mous embedded ovicell and the semicylindrical mucron inclined in the aperture. Balcombian beds at Muddy Creek, Victoria. bic. 7. Yakariavcrassocirzca:newespeciesaaace ae roe eee eee eee 24 Portion of the free, cylindrical zoarium. Janjukian beds at Bairnsdale, Victoria. Er. 8: ‘Chastosella lameliata, news species)... ae oe eee ee eee 25 Wide frond of the lamellar zoarium exhibiting a group of ovi- celled zooecia. Balcombian beds at Muddy Creek, Victoria. Fic. 9. Schizoporella orbiculifera, new SPeCieS..........0...eeeeseeecee 28 Zoarium exhibiting a perforated ovicell. Janjukian beds at Anticline Creek, Dartmoor, Victoria. NO. 9 TERTIARY BRYOZOA—CANU AND BASSLER 5I PAGE Fic. 10. Chiastosella parviporosa, new SPECiI€S............c ccc ccccee veces 2 The unilamellar zoarium composed of large zooecia with the frontal ornamented by relatively small pores. Janjukian beds at Flinders, Victoria. Pig ia. Schizoporella clypeata, NEw, SPECIES. ..6 on s4 ccs as occ cdo ssaw vanes 30 The unilamellar zoarium showing the disarranged apertures and the zooecial shield surrounded by areolar pores. Janjukian beds at Anticline Creek, Dartmoor, Victoria. PLATE 7 (All illustrations on this plate are magnified, * 20) PAGE Fic. 1. Hippomenella rarirosirata, New SpeCieS... 2.5.0... cceccevaccece 35 Fragment of a unilamellar ovicelled zoarium showing the radially ornamented ovicell with triangular area and few avicularia. Balcombian beds at Muddy Creek, Victoria. Fic. 2. Hippomenella parviporosa, new SpeCieS.............ceceeeccccaee 34 Unilamellar zoarial fragment attached to a fragment of bryozoan, showing the irregularity of position of the avicularia, the smooth frontal and the small areolar pores. Balcombian beds at Muddy Creek, Victoria. EN Gamo MRSIIIELIIUUS PONOT Adm MeWiESPECIESh temo acs cascade inca ane aes ne 37 Fragment of the unilamellar zoarium showing the place of the large, orbicular, median avicularium. Balcombian beds at Muddy Creek, Victoria. Fic. 4. Smittina (°) parwviozicellosa, new SPECIES. ..455 0.662 c0 ces oe ec eee 38 Portion of a bilamellar ovicelled zoarium exhibiting the very small ovicells and much elongated zooecia. Janjukian beds at Orbost, Gippsland, Victoria. Fic. 5. Hippomonavella acutirostris, new SPeCieS.......0...500ce eee ceeee 27 Zoarium incomplete, incrusting, showing the formation of new cells and occurrence of the median avicularium. Balcombian beds at Muddy Creek, Victoria. Bie 6. Maucronellavclongata, new Species: <02.2o he soa ce wees oo bo 38 Example showing the very long zooecia with smooth frontal, one row of small areolar pores and small mucron. Janjukian beds at Anticline Creek, Dartmoor, Victoria. Brene7= horellantuberosd, Mew SPECIES acs ete Oe aoe okie 40 Surface of the bilamellar zoarium showing the frontal tuberosities and one ovicelled zooecium. Balcombian beds at Muddy Creek, Victoria. Fic. 8. Smittinella magniporosa, new speCcieS.............. cee cece ccc eee 40 Bilamellar ovicelled zoarium, showing the large frontal pores. Balcombian beds at Muddy Creek, Victoria. Bic. 6: -Porella, cylindrorosinis, new’ Species: 4.5.22 62. dak eo. boc eke. 41 Portion of a bilamellar ovicelled frond showing the cylindrical avicularium covered by an expansion of the peristome. Janjukian beds at Bird Rock, Torquay, Victoria. 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Surface of an ovicelled zoarium exhibiting the very transverse outline of the ovicelled zooecia. 3alcombian beds at Muddy Creek, Victoria. Bice eRorellasencavata, new. SpecieSaa-- heen aan eee nee 42 A wide frond in which the avicularia are quite visible in their excavations. Janjukian beds, Anticline Creek, Dartmoor, Victoria. PLATE 8 (All illustrations on this plate are magnified, x 20, unless otherwise indicated ) PAGE Fic. 1. Porella rhomboidalis parviapertura, new variety...............06- 43 Unilamellar specimen with the characteristic small apertures. Balcombian beds at Muddy Creek, Victoria. Fic. 2. Porella rhomboidalis crassimarginata, new variety...........+...- 43 Unilamellar zoarium showing the zooecia surrounded by a broad margin. Balcombian beds at Muddy Creek, Victoria. Fic. 3. Hippomenella vermicularis, new species...........-.seeeeeeeeeee 36 Fragment of a unilamellar zoarium showing the decoration of the ovicells and the two small zooecial avicularia. Balcombian beds at Muddy Creek, Victoria. ic: 4: Heppoporetla test, new 'Species...si..c.c aeelen ere se hse cae see ae 36 Fragment of an ovicelled unilamellar zoarium. The large trans- verse aperture, six oral spines and small avicularium are shown. Balcombian beds at Muddy Creek, Victoria. 1G 5. almicellariar magna newa SPECIES seis ie ete ele seer eee 43 Fragments of the cylindrical zoarium, natural size, and an example on which the zooecia have two oral avicularia. Janjukian beds at Anticline Creek, Dartmoor, Victoria. Pic. 6. Porella operculata, tlew Species 62.2152 access ot ee eee soe eee 42 Portion of the cylindrical zoarium with calcareous opercula and showing the true form of the oral avicularium. Janjukian beds, Anticline Creek, Dartmoor, Victoria. BiGH OMI Nella magna MewaSpeCleS teenie eeeteiee eee 39 Bilamellar ovicelled zoarium showing the large dimensions of the zooecia and the very porous frontal. Janjukian beds, Anticline Creek, Dartmoor, Victoria. Fre. 8: ‘Porella: baculind, -new-SpeCieSmiacuc cos at oleae css ee ORE eee eee 41 Cylindrical zoarium with wide zooecia. Janjukian beds at Mitchell River, Bairnsdale, Victoria. Fic: 9: Phylactella chapmant,, newaspecieS:-nia.ieelieeiene cele eae ee 44 Portion of the incrusting zoarium illustrating the convex zooecial frontal, the lyrule and the recumbent ovicell. Balcombian beds at Muddy Creek, Victoria. Pic.10, ‘Costasiaiconvera. news SpeCiessaeenisni eee ee eee ee eLne 44 Cylindrical zoarium illustrating the small dimensions and the occurrence of an oral avicularium on an umbo. Balcombian beds at Muddy Creek, Victoria. NO. 9 TERTIARY BRYOZOA—-CANU AND BASSLER 53 PAGE Fic. 11. Stichoporina (?) parvicapitata, new species............... eee eeee 45 Zooecial surface of the unilamellar zoarium. Balcombian beds at Muddy Creek, Victoria. Fic. 12. Trigonopora (Metrarabdotos) vermicularis Maplestone, 1902. Surface of a narrow bilamellar zoarium received from Mr. Maple- stone as Trigonopora vermicularis. The endozooecial ovicell, form of aperture and pleurocystal frontal with lateral areolae show this to be a true Metrarabdotos. Janjukian beds, Cape Otway, Victoria. Fic. 13. Hippomenella magna, new SPeCieS............ccesccccceescccees Fragment of the unilamellar zoarium illustrating the large zooecia, the arrangement of the areolar pores and form of aperture. At the middle there is a monstrous zooecium. Janjukian beds (Torquay bore no. 1, depth of 160 feet) 15 miles south of Geelong, Victoria. PLATE 9 (All illustrations on this plate are magnified, < 20) HIGHER CthOosellavuilbosa@imewsSpeCless a -eeerieeeeae hier oie ails Surface of the bilamellar zoarium with small elliptical avicularia and some zooecia closed by a calcareous operculum. Janjukian, Aire Coastal beds, Victoria. huci25. Bathoscila jaticella.. new SpeCieS x. 34. <5 a. case olsinee nes 6 aks The bilamellar zoarium composed of very wide zooecia with several rounded areolar pores on each side. Janjukian, Aire Coastal beds, Victoria. Biers Sue eOchelia gry Gndis, NEW SPECIES. \os.c sme Aene 3 5 oetewse ores teideuele/sieiare Surface of the unilamellar zoarium with some ovicelled zooecia. One cell has a calcified operculum. Balcombian beds at Muddy Creek, Victoria. . Schizoporella tenuilamellosa, new specieS..............cceeeeeees Surface of the very thin bilamellar zoarium with a small orbicular and a transverse avicularium before each aperture. Balcombian beds at Muddy Creek, Victoria. Schizoporella macgillivrayi, new species...............cecccceees Fragment of a hollow cylindrical zoarium exhibiting the disar- ranged zooecia. Balcombian beds at Muddy Creek, Victoria. Pigs GO. i Schorotdes. erecia, NCW. SPECICS...cco lace aes oe cowie ovine ccc eae The free erect, cylindrical zoarium composed of large elongated zooeCcia. Janjukian beds, Gellibrand, Victoria. IGS. 7; 6. Jidymosella- clypeata, new: Species. . 2 tui... i sie se niece sees: 7. Cellular side of the free, unilamellar zoarium. The frontal shield, with its two foramina, is large and smooth. 8. Dorsal side of the same specimen. Balcombian beds at Muddy Creek, Victoria. Fic. _ Fic. oe 35 32 29 29 33 S4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 93 Fic. 9.° Schizoporella arcana, new: SPeCieS. 0 6i..6 suse sin nen se eis visas oie 30 The unilamellar zoarium showing the large perforation in front of the aperture. Balcombian beds at Muddy Creek, Victoria. Fic. 10. Microporetla cailleti, new Species’ 0.02. os acres cis laminates a The incrusting zoarium with several ovicelled zooecia. Balcombian beds at Muddy Creek, Victoria. Fice1i. Fenestrulina practexta, tlew Species... 74h a. es ee tay eee 34 Surface of the bilamelar zoarium showing ascopore much re- moved from the aperture. Janjukian beds at Flinders, Victoria. SMITHSONIAN MISCELLANEOUS COLLECTIONS MOVES E)si5 INO Sa tebe 1 TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA (For explanation, see page 45.) MOS Ey tes 2 VOL. 93, 46.) planation, see page (For ex = _ < ig kK 17) 3) < < 4 O FE 2 > = O ig w < O N O > i it W 2 O FE 0) O = WW L O > © S k © WW kb SMITHSONIAN MISCELLANEOUS COLLECTIONS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES 9/3) NOS 9) PEs TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA (For explanation, see page 47.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. ‘9135 NOA*9), (PL 4 TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA (For explanation, see page 48.) VOL. 93, NO. AUSTRALIA , 49.) page see (For explanation, = x O FE Z > = (9) iN WL < O N fe) > [V4 o Ww = O E 1) O = Wl a6 O > Na = kK V4 WW kK SMITHSONIAN MISCELLANEOUS COLLECTIONS SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.-93), NOn9, PEG TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA (For explanation, see page 50.) VOE 93) NO=.9)) PIE: 7 SMITHSONIAN MISCELLANEOUS COLLECTIONS . Es esas. ee en . TERTIARY CHEILOSTOME BRYOZOA FROM VICTORIA, AUSTRALIA ‘xplanation, (For S CTION MISCELLANEOUS COLL A\THSONIAN SM s J < '6 = n 2) < < r i) Fk ¥ > = O '4 re < O N ce) > r 0 Wu = O Ff 0) ce) = WW ae O > 4 < F © W = ) 53 see page (For explanation, SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 93, NUMBER 10 (End of Volume) AN INTRODUCTION TO NEBRASKA ARCHEOLOGY (WiTH 25 PLATES) BY WILLIAM DUNCAN STRONG Anthropologist, Bureau of American Ethnology (PUBLICATION 3303) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 20, 1935 The Lord Waftimore Mreas BALTIMORE, MD., U. 8 A, CONTENTS PAGE HORE WOU Waitrose oscar catnist ole: aieret pene sneavatenete TNS PE Pe es are coats Sire rosap rere shoe I MTG OCIS Hl ONT Meee eeed ce sheto tc ehalcvoner cha creek crete lome euniee wich eed stereu suas veya are) sesnaudialede cy shay 6 Eistoniceanautracitional backeround es acccimee eclceciee elsre ee irra. «aes 8 Table 1, a digest of certain maps showing the Indian tribes in the region Ge ehrasica, GEOM TOFS 10 TS1Qscn nic dee anise ce a cor ssa lt ara Oia) ane aie 14 RHEE VATOMMIetItal SELEMAR. 5/25 c> ccc ade eles ese naira das dle «ia eleaiwajoeeis’ 30 Previous archeolosical work in Nebraska soc 622 ssc ce cece cect een nce es 40 Historic and protohistoric Pawnee culture as revealed by archeology...... 55 Prehistoric sites recently excavated in Nebraska.....................5. 68 Village on Lost Creek (Dooley site), Franklin County.............. 69 Methods#otvearth-lodgevexcavationesracten scree siecle arm ere 73 TESTS AC PAS OP See Me Re eae A Ost ade Leo 5s 1 eet 74 StonacewGu Caches pits 1. aoe a eaey ears cia tie WeiheGa solace as © 78 IRELUSGush Aap Siitat state a ookrteeenst aes erate la rscs ane, mike veteecuere eetarsinys 8I REN BES) sy ste eae PIE SON oe a ear ORAS ROLE eg Arcee ea 82 Wroslcsint around -Stone.....cos as sre ci ore eos esi da cient saa 85 Wotkeintchipped sstomel ck cinetien er crc a oen eccrine ele aicrerats 87 Table 2, classification chart for chipped points........... 89 Table 3, distribution of chipped points according to type.. 90 VG Teka EN AD TNS pecs ed stata su aci-a) ober eere a reer oan eaten oreo tar arabe hers 94 Bonesonrantler bracelet tragmentse- eee oe cine een 97 Wor kmeittica mt lemyasesterrsroseraiscewe tokercccie aie caste thee ses tensin tactics ous ieie oie 97 WViorkeai ties helllimyamraystters «ecg wetter welt eat eRe on Pada s orc yes eaete 100 Animal sandemolluscan remainsaaseeee ee enero eee eriion. 100 PV OUS Chee rer eecacope cette ctatek acest vote etc crore ae ee Nace ora iste Locicre reiave aie IOI Disposal wotmethexd ead -r4s:>.ccc anes cae ete OL ASL eae 103 Ossuary on Prairie Dog Creek (Graham site), Harlan County....... 103 IPieyineray ainoyan’ (Grmneyan CRSVEIAY. boc nocaudacanononnaacnaoeneac 108 Worle inisekOundsStOne sans ast eames ier eeterenicale ts Gees 109 Wronkeitiachippnedeestoner scarp rraccrart relate atoice tera cunsie chore 109 ‘WACGiel eathat jl Och ates ees eenesterenenctene Siege ian 4 oer cine eee ee eee eee IIO Wis clteyhnl Ghintd Lene pista oenctoso 6 Goss es 6 OU8.A6 DIO RTOCIs Set ORI nE ee aE III \iWferel Feorio ats) oe) iieepencane cee tera. ctod cic ort ols cao) ec caer Ona aera een III WOE SIN COPPER wut sorsencic ert reyerertehe release cacy cuaye, ies e e| Slo teases 114 Negative excavations in vicinity of Graham ossuary.............. 114 Conclusions regarding Prairie Dog Creek sites.................. 116 Ossuary on the Republican River (Marshall site), Harlan County.... 116 AGutactse frome Viacsialleosstiaisyereeem neces tenis seca ccccie oes 120 Gonclusions regarding Marshall .ossuatry.. ....... 0... 20 ens ce eee 121 Ossuary on Republican River near Alma, Harlan County............. 122 Ossuary, on) Munson Creek, Eloward! Gountya... 20. s5es5606 500-0000 123 1V CONTENTS PAGE Village near Rock Bluffs (Rock Bluffs site), Cass County............ 124 1a lalb (en Celene eRe eRe cor ant mine Cin aa cncbiaca co oapedwcatc 126 TOUS EF 2 sg hi seelais ari Gan phi roe Mee arene RTECS ORS reac eee 126 Storage: spits: 2vsn dure cowee eae oer e ee eee te 129 Refuse heaps: asd sicisietommatay teerneasrncecte eres eae RO Ore eR er ar 130 Pottery irom SHOuUse 2... 4 ees Shel en eee ao Tee ieee 130 Work: inground) stone: ccpenacestee ae eee eee ee eel 133 Work im chippedestone: cn. ssacect oc croeinsor rene niece emote 134 Wrork “inebone’ssertc.s cele Se eye orn eenatercelorccsicce aC ener eee 136 Weorkearp. sShrelisc; crise scsessahe tune oraercianttne Decne ene SO ner 136 Baishketiteyys aa on-vacherithorsntyal toa ersvestadehe bccew ER OSes TEE ho OT 137 Animal sancdenolliascane nema creer ene taeieicre 137 TOUS Es 3 ecdey storie ase aR CIR OTS TOO OO re Orion 138 Disposalvot the dead’ 44 ciranttesc marcus esener Oe eee Tete 139 laltoyb Re Suite saree he INGerelte leiouniky (Cave S))oca0dcunccucecadouencsoocr 139 IPOETE tay? wanes evapo s ci egciove crapoicepotcesyere el ner OTSR IPAS ere eran 139 Workin eround, Stoney <:s2.scccathee sa urls Serecsie cies euren tater 140 Wonk? in ‘chipped istome ap cicversa custeccsiere casveraterachscree treteretene eteetete 142 Workin bones scvcocn bismnocie acne Cie ee aeata ace te) Sees Cer OTe eee 142 Vegetal (remains? < .g nec wine coucsaye sce doris Oe ee eres Reno 143 Village between the Platte River and Papillion Creek (Gates site), Sarpy County; jscet ca coc eae ates cure oie sient toe Coach wieepee erate iene mas 143 | Fob oy si Cae tie 7 wee OID ES