liiiiliKjiiHiliilJlliJiiiiiiPPiiiliiiiiiiiiii W^ ■i^iilliil^iiil' i4 ^>u\ M 'ilijijlijiii ^illi n PROCEEDINGS OF THE AMERICAN ACADEMY OF ARTS AND SCIENCES. Vol. LVII. FROM MAY 1921, TO MAY 1922. BOSTON: PUBLISHED BY THE ACADEMY. 1922. The Cosmos Press cambridge. mass. CONTENTS. Page. I. The Grid Structure in Echelon Spectrum Lines. By N. A. Kent AND L. B. Taylor 1 II. The General Conditions of Validity of the Principle of Le Chatelier. By a. J. LoTKA 19 III. The Effect of Tension on the Electrical Resistance of Certain Ab- normal Metals. By P. W. Bridgman 39 IV. Notes on the Early Evolution of the Reflector. By Louis Bell . . 67 V. The Effect of Pressure on the Thermal Conductivity of Metals. By P. W. Bridgman 75 VI. The Failure of Ohm's Law in Gold and Silver at High Current Densities. By P. W. Bridgman 129 VII. A Table and Method of Computation of Electric Wave Propagation, Transmission Line Phenomena, Optical Refraction, and Inverse Hyperbolic Functions of a Comple.v Variable. By G. W. Pierce 173 VIII. Artificial Electric Lines with Mutual Inductance between Adjacent Series Elements. By G. W. Pierce 193 IX. The Parasitic Worms of the Animals of Bermuda. I. Trematodes. By F. D. Barker 213 X. Additions to the Hydroid Fauna of the Bermudas. By Rudolf Bennitt 239 XI. Some H ymenopterou^ Parasites of Lignicolou^ Itonididae. By C. T. Brues 261 XII. A Revision of the Endogoneae. By Roland Thaxteu .... 289 XIII. The Echinoderms of the Challenger Bank, Bermuda. By H. L. Clark 351 XIV. Atmospheric Attenuation of Ultra-Violet Light. By E. R. Schaeffer 363 XV. The Ratio of the Calorie at 73° to that at 20°. By Arnold Rom- berg 375 XVI. Studies on I Jisect Spermatogenesis. IV. The Phenomenon of Poly- megaly in the Sperm Cells of the Family Pentatomidce. By R. H. BowEN 388 CONTENTS. IV XVII. Note on Two Remarkable Ascomycetes. By Roland Thaxter. 423 XVIII. Records of Meetings 437 Biographical Notices 470 Officers and Committees for 1922-23 623 List of Fellows and Foreign Honorary Members 525 Statutes and Standing Votes 545 RuMFORD Premium 561 Index .... 563 57-1 Proceedings of the American Academy of Arts and Sciences. Vol. 57. No. 1, — December, 1921. THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. By Norton A. Kent and Lucien B. Taylor. Investigations on Ligbt and Heat made and publishgd with aid from the RuMFOBD Fund. (Continued from page 3 of cover.) VOLUME 57. 1. Kent, Norton A. and Taylor, Ltjcien B. — The Grid Structure in Echelon Spectrum Lines, pp. 1-18. December, 1921. $.75. Proceedings of the American Academy of Arts and Sciences. Vol. 57. No. 1. — December, 1921. THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. By Norton A. Kent and Lucien B. Taylor. Investigations on Light and Heat made and published with aid from the RuMFORD Fund. THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. Norton A. Kent and Lucien B. Taylor. Received July 7, 1921. Presented October 19, 1921. Some years ago Nutting ^ noted a peculiar, complex structure, termed by him the "fluting" or "grid," which appeared in many echelon spectrum lines, and consisted of several fine components of different and often changing intensity. Later one of us ^ independ- ently noted this structure. Nutting crossed the 12" Lummer plate of the Bureau of Standards with his echelon and was apparently forced to the conclusion that the structure was real — that is, that it indicated an actual discontinuity of emission in the source. Proceeding on the assumption of reality, the writers attempted a solution of the problem using LiX 6104 which, although known to be a spectroscopic doublet, offered peculiar advantages in that the grid was extremely brilliant, well-marked and persistent. Apparatus. The apparatus used consisted of: — Two echelons: No. 1, made by Porter, 30 plates, each 14.76 mm. thick, step 1 mm., aperture 31.0 by 33.0 mm.; No. 2, made by Petit- didier, 30 plates, each 23.29 mm. thick, step 1 mm., aperture 31.0 by 35.5 mm. The Bureau of Standards 12" Lummer plate kindly loaned by Dr. Stratton. A Hilger Lummer plate — length 131 mm., width 14.5 mm., depth 4.827 mm. A Hilger constant deviation prism spectroscope combined with an echelon as in Figure la; also a separate Hilger spectroscope with another echelon spectroscope as in Figure lb. The achromatic lenses of both echelon spectroscopes are of about 50 cm. focal length and 5 cm. 1 Astrophys. Jour. 23, pp. 64 and 220. 1906. 2 Kent, Proc. Am. Acad. XLVIII, No. 5. Aug. 1912. KENT AND TAYLOR. aperture; each echelon bed rotates on an axis at its center; the Hilger micrometer is fitted with one fixed and two movable cross-hairs as shown in Figure 2. ^J7> Fig. lb. Figures la and lb. S, slit; L, L, lenses; E, echelon; P, prism; O, ocular. Figure 2. SS', fixed crosshair; MM', MM' movable system; LL', spectrum line. A Littrow mount spectroscope consisting of a Petitdidier achromat — focal length 30 feet, aperture 6"; and an An- derson grating — aperture 3f " vertical by 5" horizontal, 15,000 lines per inch, used in the third order. A 5 K. W. transformer, 1 10 to 30,000 volts ratio of trans- formation, fed by a 60 cycle Holtzer-Cabot 4.5 K. W. gen- erator. Various large induction coils capable of giving 6" sparks and operated by a rotary mercury break and an electrolytic interrupter, had proven insufficient. A vacuum arc of construction as indicated in Figure 3. THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. Figures. One fourth original size. D, D, fibre disks; O, O, water outlet; P, P, fibre plugs; A, A, asbestos; J, water jacket; W, W, W, windows; I, I, I, water inlet. 6 KENT AND TAYLOR. This was also adapted to pressures of several atmospheres as the glass windows and fibre plugs were held in place by threaded rings. Quartz vacuum tubes — even pyrex glass having proven unsuit- Water infahe i ^^ 2;z^ ry 7Z2L I To vacuum pump \ Wafer ouiki "Wafer /'ntaKe I i -rrj « ••:•■•'•■-: ■v_ 1 Waier ouilet Figure 4. C, C, cork stoppers; R, 11, rubber sponges; T, T, terminals. T To vacuum pumps Tc H, generaior and dryino apparatus Figure 5. T, tube; M, manometer; B, bulb. able — of various forms, the most successful of which, for salts such as lithium chloride, proved to be that shown in Figure 4 in which fine brass wire, often in helical form, was fitted into brass caps, 6 mm. in THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. 7 diameter, and sealed in with De Khotinsky cement, each joint being cooled by a water jacket. The salt is shoved into the capillary by a wire and the tube will run many hours without refilling. It may be used end on as well as side on. The capillaries varied from 2 to 0.5 mm. Auxiliary apparatus as shown schematically in Figure 5. The bulb B, prevented too rapid changes in pressure. The system was washed out with hydrogen from a Kipp generator, dried by sulphuric acid and a calcium chloride tower. The mercury manometer, IVI, indicated the pressure — generally from 8 cm. to a fraction of a millimeter. Procedure and Certain Results. Both Lummer plates were each in succession crossed with echelon No. 1. In each case, with a carbon arc soaked with lithium chloride, both at atmospheric pressure and in a moderate vacuum, there appeared a pattern which, at this stage of the investigation, seemed to indicate that the grid was real. The following facts, (1) to (6), are, however, clearly not in accord with this conclusion, and prove con- clusively that this curious structure is due to the phenomenon of "secondary maxima" observed by Stansfield ^ and resulting from successive reflections from the surfaces of the echelon plates, producing a Fabry and Perot system in the region of the primary light of the echelon. (1) to (3) deal with some of the criteria of echelon secondary maxima given by Stansfield. These criteria are, in essence, indicated below by italics. (1) The width of LiX 6104, given by an open carbon arc at atmos- pheric pressure, as seen in the Littrow grating, using a narrow slit, was found to be about 0.25 t. m. when echelon No. 2 showed the grid plainly. The suspicion, therefore, was confirmed that the line was too wide for the echelon, the difference between the adjacent orders being about 0.26 t. m. In the case of Janicki's observation * of Hg. X 5461, Nutting's work on lines of many elements, and the work of one of us on the zinc lines as given by arc and spark, the indications are that with all lines for which the echelon shows the grid, their breadth is so great that the use of this instrument is not at all justifi- able. The writers then proceeded to study the structure from this new 3 Phil. Mag. (6) 18. .383. 1909. * An. der Phys. Vol. XIX, p. 36. 1906. 8 KENT AND TAYLOR. standpoint, considering the primary line of width approximately 0.25 t. m., and not as formerly, one of the grid components itself. These componcnis are indeed, in this sense, each narrower thari the primary maximum — 0.25 t. m. — the grid components, all of them now regarded as secondary maxima, being only about 0.05 t. m. in width in echelon No. 2. (2) The curvature of one of the mercury yellow lines was compared with that of a^ grid component in X6104. By stopping down the echelon spectroscope slit, a line of definite length was observed, and by setting the stationary cross-hair of the filar micrometer upon the ends of the image, and the movable system upon its center, the horizontal distance, d. Figure 2, from the ends of each line to its center were measured. It was found that the curvature of the com- ponent is about 25% greater than that of the primary line. (3) With a small mirror, set at 45°, over the lower half of the echelon spectroscope slit an argon vacuum tube and the lithium arc were observed at the same time. The relative motion of the grid compo- nents in X6104 and a nearby argon line were then studied as the echelon was rotated. The primary argon line moves about one-half as fast as the grid components. Quantitative measurements of the relative displacements were later made with Zn X4810. A quartz vacuum tube was fitted with coiled brass wire leads and brass terminals, exhausted, filled with hydrogen to 10 or more cm. pressure and then gradually exhausted to 1 mm. or less. The zinc lines given by the brass wire leads appeared very sharp, steady and brilliant. With X4810, as thus produced, was com- pared the "gridded" line of a cored carbon arc at atmospheric pres- sure, in which small pieces of zinc had been placed, the small mirror arrangement allowing simultaneous observation of both sources. Upon rotation of the echelon the grid components rushed by the narrow tube line. To measure the relative speed a plane mirror was attached to a side of the echelon case. The image of an illuminated slit in a piece of cardboard was formed by a lens upon a distant scale after reflection from the mirror. The echelon was set near the ^ = 0 position. A reading of the position of the slit image on the scale was taken when the tube line lay upon the fixed hair of the filar micrometer. The echelon was then rotated until the slit image moved about 2 cm. The displacement of the tube line was then measured by the movable cross-hair system. A similar series was then taken with a grid line. The ratio of the displacements was 3.6 : 6.4 or about 1 : 2. (4) The echelon was removed and the ocular focussed on the prism THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. 9 image of a line. Replacing the echelon shortened the focus for a true narrow echelon image by about 0.6 mm. The focus for the grid com- ponents of the same line was 0.7 mm. shorter yet — the light forming the grid had traversed the echelon plates more than once. (5) Although the grid components are generally very well defined (the minimum being " deep"), it is a difficult matter, with a fluctuating source such as an open arc, to obtain accurate measurements. The grid spacings appear to vary slightly at different stages. When the grid is complete the spacing is regular and, within the limits of error of measurement is equal to one fifth the distance between the orders. This was proven as follows : — A quartz tube having merely coils of fine brass wire as terminals gave extremely fine zinc lines. Echelon No. 2 was set in double order condition and Ao, the difference between the two orders, measured for X4S10 (see Table I). Then the grid was measured as given by a 3 ampere open carbon arc. Three distinct series of readings were taken. Then the tube was again used. The accuracy of an individual setting was about 0.2% in Ao and about 5% in Ag. It thus appears that in this region, at least within 2%, Ao = 5Ag. The focus of the instrument was, of course, not changed, the dift'erence between that for primary and secondary maxima being so slight that distances between the components of the grid are not appreciably affected. TABLE I. Distances are measured in divisions of the|micrometer head. Each Ao dis- tance given is the mean as calculated from four settings; each Ag f rom two. Settings were made on the six centrally situated grid components. Ao for Zn X 4810 Ag for six grid components. 22.60 22.50 Mean 22.58 1-2 2-3 3-4 4-5 5-6 Mean Mean of Means 4.6 4.3 4.8 4.3 4.8 4.4 5.0 4.6 4.6 5.0 4.8 4.8 4.3 4.7 4.1 4.7 4.6 4.5 4.6 22 58 -^ 4 . 6 = 4 . 9 or Ao = 4 . 9 Ag A similar series for Zn X6362 gave Ao = 27.65 and Ag = 5.6, 5.9, 5.5, 5.6, 5.4: mean = 5.5. Hence Ao = 5.0jAg. 10 KENT AND TAYLOR. (6) The structure given by both echelons is the same. That is, there are five secondary maxima for every primary maximum. Ag for Hydrogen X6563 is 0.061 t. m. for instrument No. 2, while for No. 1 it is about 0.09G t. m., which again is another fact fatally inconsistent with the existence of a definite discontinuous emission in the source. With this evidence at hand the writers then attempted to clear up the results of the crossed dispersions. The source previously used was hardly adequate. By removing the soft core of the lower carbon it was possible to feed copiously into the arc a strong LiCl solution. Greater brilliancy and steadiness were obtained. The results were unmistakably in accord with the facts given in (1) to (6) above. Figures 6a and 6b indicate the structure observed. These will be y 8' o \ <■■' yp P V Fig. 6a. Fig. 6b. Figure 6a. Li X6104 with crossed Lummer plate and echelon. Grid not indicated. Xi and X2 in sing;le and double order condition respectively. i Figure 6b. Li X6104 as in Fig. 6a. Grid shown. Xi and X2 both between isingle and double order condition. discussed in full below (see page 16). When one component of the spectroscopic doublet is in double and the other in single order condi- tion three lines appear; when both are in a condition between single THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. 11 and double order there are four lines. It is probable that these four lines, under conditions of inferior illumination, were interpreted as four separate and true lines. It is unfortunate that at first the only line available for study was a doublet. With this latter and better source a zinc chloride solution gave X4810 sufficiently strong. The crossed dispersions prove it to be a simple, though broad, single line when the echelon alone shows the grid. Further Results: Characteristics of the Grid. (a) From numerous observations upon Li X6104 and Zn X4810, as developed by various sources, such as vacuum tubes and arcs (on 110 and 220 volt D.C. circuits and from 1 to 20 amperes) under high, normal and low pressure, in which the cross-hairs of the filar micro- meter were set successively upon the true, narrow, lines given by the tube and the grid components given by the arc, it is quite certain that the grid is built up approximately as follows: — Suppose that in a hypothetical grating of resolving power and dispersion equal to that of the echelon, a line which is at first very narrow, e.g., 0.025 t. m., gradually becomes less monochromatic, owing to changing conditions in the source, and appears as represented diagrammatically by the small letters a to e, Figure 7. Four cases must be discussed as shown in Figures 7 to 10, respectively. Case I: — The echelon in double order condition gives successively images A to E. When the line is very narrow the echelon shows it as such, in A. Similarly for a line of width, Ag — the width of a grid component or an intergrid distance — it is shown as in B. When of width 3Ag, the echelon shows no change, C appearing as B; for, at m, the primary and secondary action together give a decided minimum. When the line has a width, as in d, the echelon shows a triplet, D, and when of width as in c, or greater, the grid is complete — five grid maxima, 1 to 5 and 6 to 10, for each maximum, such as 3 and 8, which a narrow line would give; four maxima, 4 to 7, between the double order positions, 3 and 8, of such a narrow line. For a given position of the echelon these grid components do not, in forming, move very much, if at all: they come up in situ. There exists an apparent motion, in and out, which is probably due to the changing width of the primary line, which may not at all times be such as to complete the entire width of a grid component. Case II: — When the position of the echelon, its temperature and the wave length of the line observed, result in the central grid minimum 12 KENT AND TAYLOR. I a A ^.. 1 n ^^ b B in '% i C -JAX- 1 E ^ $: 8 ' 1 1 D ^7AS— 1 ^ ^ ^ e ^ % 1 H !■ / 2 3 V 5 6 7 3 5/0 Fig. 8. Fig. 9. Fig. 7. Figure 7. Case I: Echelon in double order condition and a grid maximum coincident with the primary maximum. Figure 8. Case II: Echelon in double order condition and a grid minimum coincident with the primary maximum. Figure 9. Case III: Echelon in single order condition and a grid maximum coincident with the primary maximum. , THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. 13 occurring in the position of the narrow tube line, as in Figure 8, for a double order condition of the echelon, there are nine or even eleven components when the grid is strong. Note that the grid components 2, 3, 7, 8, which at first are very brilliant when the grid is "young," grow weaker, 2 and 8 often being so faint that it is difficult to make accurate micrometer settings upon them. Case III: — The treatment is the same for a single order condition of the echelon, as in Figure 9 which shows a triplet, quintuplet, or, with neighboring parts of adjacent orders, even as many as eleven components. Case IV: — Here a grid minimum coincides with the primary maximum and the grid com- ponents are as shown in Figure 10. The above statements explain why an origi- nally narrow line, as its width increases, may ap- pear, as it actually does, a triplet or quintuplet, as in Figures 7 and 9, or may, as it were, "reverse" and then quadruple, as in Figures 8 and 10. Actual reversal as shown by the grating probably occurs much later in the historv of the line. (See page 15.) Further, if a line be intrinsically unsymmetri- cal, shading off to the red for instance, the sec- ondary action masks an early stage of broadening, and the left grid line, 2, forms as in A, Figure 11. Line 3, as in A', then comes up as 2 strengthens. (b) The grid begins to disappear and the line gradually becomes broad and structureless when the primary line exceeds 2Ao in width, Ao being the distance between two adjacent orders. This was determined as follows : — Using as narrow a slit as possible, a low power ocular and a mm. scale, an eye estimate was made of the breadths of various portions of an arc line shown by the grating. These were reduced to t. m. The same source was viewed simultane- Fic. 10. Figure 10. Case IV: Echelon in single order condition and a grid minimum coincident witii the primary maximum. 14 KENT AND TAYLOR. ously by echelon No. 2. For Zn X4810 three components of the grid exist when the grating shows a line 0.12 t. m. broad. Ao for X4810 = 0.155 t. m. f X 0.155 = 0.09 t. m. which compares favorably with 0.12 t. m. The complete grid exists when the line is 0.3 t. m. or 2Ao broad and the image begins to pass into a structureless line at 3Ao. Similarly for Li X6104 a full and well-marked grid exists at a line width about 0.2 to 0.5 t. m. or Ao to 2Ao (as here Ao = 0.25 t. m.). The grid is poorly marked above about 2Ao and is gone at 3Ao. (c) Numerous lines in the spectra of Na, Hg, Fe, Mg, Cd, Ca, Sn, Pb, and Bi, developed by an open carbon arc, show the grid whenever the line is sufficiently broad — rendered so by introducing more of the substance or increasing the current; also by increasing the capac- ity in the case of a spark. \d) LiXXG708 and 6104, Zn XX4810, 4722 and 4680, also HgX5461 (mercury being fed into the lower cored carbon) show by their behavior that a line which is too broad will appear structureless in the echelon, that the center of the core of an arc may show the grid complete while light from the wings of the image gives a simple structure of but one to three components. With a sufficient amount of vapor the com- plete grid may be obtained even at low pressure. (e) A study of Zn X4810, from an arc in the vacuum or pressure tank, at pressures from 2 cm. of mercury to about three atmospheres, showed that moderate changes of pressure do not produce measurable displacements in the grid components, but merely alter somewhat their relative intensities, shifting the maximum over one or two com- ponents or even bringing up new ones. This of course means that, as long as a grid exists, the components do not change appreciably their position with changes of wavelength as small as 0.015 or 0.020 t. m.^ Their position is affected more strongly by the position of the echelon and its temperature. Similarly, the grid components of the spectroscopic doublets Li XX6708 and 6104 developed in vacuum tubes show intensity shifts with changes of pressure over the range of one atmosphere. (f) The " end on " position of a vacuum tube will generally show a more complete grid than that "side on." (g) If a line broaden unsymmetrically with increase of current the maximum of intensity will shift. Those components which are just being formed show an apparent motion outward as the number of 5 According to Humphrey's and Mohler's results for Zn, the pressure shift reduced to X 4000 is 0.057 t. m. for twelve atmospheres. THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. 15 components increases, the first step resembling a narrow reversal as in Figures 8 and 10 or a central fixed line with two moving wings as in Figures 7 and 9. But the writers feel that this apparent motion is due to the fact that each grid component is not formed in ioio at once: the part which lies nearest the center of the system is formed first. Certain it is that this apparent motion ceases abruptly when the component has reached a position which is one grid distance from its neighbor. If the source be an arc, many rapid fluctuations in intensity occur. (h) Although the resolving power of the grating (225,000 in the third order) is far below that of the echelon (about 750,000 for X6100 for echelon No. 2) it is hard to reconcile the images given by the two instruments on any other assumption than that the grid is due to secondary action. To throw further light on the problem, Li X6104, given by a vertical carbon arc soaked with LiCl, was viewed simultaneously by echelon and grating. Table II gives a summary obtained from various ar- rangements. TABLE II. 1 1 1 1 indicates the grid; ■ a broad structureless line; | a narrow unreversed line, or one very slightly reversed; || a broad and strongly reversed line. Arrangement Sign of upper pole IVile soaked with solution Echelon shows Grating shows 1 + + At + pole nil « _ « H II 2 + " + " nil " - " ■ II 3 — " - " ■ " + " nil II 4 + « _ « B " + " nil II 1 Therefore which pole is soaked makes no difference, nor does it matter which pole is above. The region near the + pole generally shows the grid in the echelon, that near the — pole a broad structure- less line. The grating always gives a narrow unreversed line or one very slightly reversed where the echelon shows the grid, and a strongly reversed line where the echelon shows no structure. Thus the grid does not result from conditions which produce a reversed grating line. 16 KENT AND TAYLOR. With Li X6708, which usually appears widely reversed in the grat- ing, the grid is more difficult to obtain in the echelon, while with Na X4972 — given as an unreversed line by the grating at either edge or centre of the arc image — the echelon shows the grid at both edge and centre. (i) We are now in a position to discuss jn detail Figures 6a and 6b. These were obtained with the 131 mm. Lummer plate set between the collimator and prism of Figure lb and crossed with echelon No. 2. The source was that described on page 10: the arc current being from 10 to 25 amps. The plate dispersed vertically, the echelon horizon- tally. Both figures are drawings based on visual filar micrometer measurements, a single cross hair being moved successively along the axes, vv' (vertical), hh' (horizontal), aa' (across the structure) pp' (parallel to it), as shown below the two figures. Two Lummer plate orders are shown in each figure, the primes distinguishing these. The numerals indicate the two components of the spectroscopic doublet, the breadth along axis aa' their approximate relative intensity. Xi is the weaker line, Xo the stronger in both figures — X2 being the component of longer wavelength. In Figure 6a X2 is in double order condition; in 6b both Xi and X2 are between double and single order. The echelon grid structure is not indicated in Figure 6a: in 6b its approximate position is shown. It was difficult to observe at the ends of the lines and so is not there indicated: it is slanted at an angle of about 2.4° (see gg' in Figure 6b) with the vertical. The slant of the lines themselves as well as that of the grid changes with the positions of both plate and echelon: further, the grid slant is not due to the curvature of the echelon image. This may throw some 'light on the disappearance of the grid at a breadth of line greater than 2Ao. For, as the echelon action alone is given by the projection, on the pp' axis, of the grids of the lines Xi and X2, it is evident that lack of coincidence owing to slant would tend to obliter- ate the grid altogether, this indicating that two broad lines, the centers of which lie as far as 0.1 t. m. apart (the AX of the two components of Li X6104), may not give coincident grid structures; or, in other words, the grid maxima do not (for any one position and temperature of the echelon) necessarily fall together. This is not inconsistent with shift of intensity for small changes of wavelength (0.015 to 0.020 t. m.) as noted on page 14. Shift of intensity and position probably both enter with change of wavelength of the center of gravity of a primary echelon image. These two figures show that the grid is unquestionably a secondary THE GRID STRUCTURE IN ECHELON SPECTRUM LINES. 17 echelon action. Otherwise the regions between hnes 1 and 2 would have been filled in with a structure along axis aa' similar to that along pp'. With an echelon alone we have obtained only the weaker component of Li X6104 as a single narrow line. We plan to cool the tube with liquid air, thus sharpening the stronger component so that it will no longer suffer the secondary action, to which the small satellite is probably due. (j) We have no record of having observed in either echelon any ungridded line of width greater than Ag. Either there exists (1) a very narrow line, (2) an irregular series of such, as, for instance, in the yellow mercury lines, (3) a line of width Ag, (4) a series of such (the grid more or less complete) or (5) a broad, structureless image cover- ing between one and two orders. And it appears extremely probable that the "reversal" of the main component of HgX5461, noted under certain conditions by several observers and often noticed by us, may be modified by the entrance of secondary action due to the excessive breadth of this component. (k) The retardation producing the primary maxima of a narrow line is proportional to 7i — l, while that of the light undergoing secondary action is proportional to 3w — 1. Thus the difference in retardation in case of the two actions bears the ratio to the retardation of the 2n . . 2w primary of ;, which is a function of n alone. The value of r n— 1 n — I varies from 5.50 for X6563 to 5.37 for X4341 in echelon No. 2; and from 5.48 to 5.35 respectively in No. 1. Since echelons are generally made of substantially the same kind of glass, any two having equal separation of primary orders will have equal separation of secondary maxima, because this separation is the same fractional part of the separation of the orders; but the values of Ag in t. m., varying with the dispersion, will, of course, differ in different instruments. We cannot state just why Ao = 5Ag. The measurements given above indicate that this is so within the limits of experimental error for both the violet and red regions. It would be interesting to assemble an echelon under water, press the plates together and allow the superfluous water to drain off. This process might vastly reduce the secondary action. If successful Canada balsam might be substituted for water thus producing a more permanent instrument. We plan to try this experiment shortly. 18 KENT AND TAYLOR. Conclusion. Summarizing the above results, we may state that the evidence is entirely against the existence of a discontinuity of emission in the source. The grid is due to a secondary action of the echelon which enters when the line under investigation is not sufficiently monochro- matic. This means that the previous work of one of us ^ must be considered as of small value and also that ^n explanation of the apparent complexity of structure obtained by Nutting ^ can be found in secondar}^ action. The results obtained emphasize the fact that when an echelon is used to measure small wavelength differences, great care must be taken to obtain the lines so narrow that their width is less than i Ao, else secondary action may enter to cut off an edge of a line and thus give a false intensity-maximum position. We must record our appreciation of the help rendered by various student assistants, especially Alessrs. Greenleaf and Risga. We are also indebted to Dr. Lucy Wilson for her skilful aid during part of this research and to our assistants. Miss Pearson for mathematical work in connection with the calculation of the constants of the echelons, and Mr. Gilman for making the sketches accompanying this article. We wish also to thank sincerely the Rumford Committee of the American Academy for numerous grants which made possible the purchase of the main pieces of apparatus used in this investigation. 6 Proc. Am. Acad., Vol. XLVIII, No. 5. Aug. 1912. 7 Astrophys. Jour., XXIII, pp. 64 and 220. 1906. Physical Laboratory, Boston University, May, 1921. VOLUME 56. 1. Kennelly, a. E., and Kurokawa, K. — Acoustic Impedance and its Measurement. pp. 1^2. February, 1921. $1.25. 2. Bell, Louis. — Ghosts and Oculars, pp. 43-58. February, 1921. $.85. 3. Bridgman, p. W. — Electrical Resistance under Pressure, including certain liquid Metals, pp. 59-1.54. February, 1921. $1.25. 4. 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PROCEEDINGS. Vols 1-56, $5 each. Discount to booksellers 25%; to Fellows 50%, or for whole sets 60%. The individual articles may be obtained separately. A price list of recent articles is printed on the inside pages of the cover of the Proceedings. Complete Works of Count Rumford. 4 vols., $5.00 each. Memoir of Sir Benjamin Thompson, Count Rumford, with Notices of his Daughter. By George E. Ellis. $5.00. Complete sets of the Life and Works of Rumford. 5 vols., $25.00; to Fellows, $5.00. For sale at the Library of The American Academy of x\rts and Sciences, 28 Newbury Street, Boston, Massachusetts. 57- 2 Proceedings of the American Academy of Arts and Sciences. Vol. 57. No. 2.— Januaby, 1922. THE GENERAL CONDITIONS OF VALIDITY OF THE PRINCIPLE OF LE CHATELIER. By Alfred J. Lotka. (Continued from page 3 of cover.) VOLUME 57. 1. K.ENT, Norton A. 6u»d Taylor, Lucien B. — The Grid Structure in Eichelon Spectrum Lines, pp. 1-18. December, 1921. $.75. 2. LoTKA, Alfred J. — The General Conditions of Validity of the Principle of Le Gbatelier. pp. 19-37. January, 1922. $.75. Proceedings of the American Academy of Arts and Sciences. Vol. 57. No. 2.— .January, 1922. THE GENERAL CONDITIONS OF VALIDITY OF THE PRINCIPLE OF LE CHATELIER. By Alfred J. Lotka. THE GENERAL CONDITIONS OF VALIDITY OF THE PRINCIPLE OF LE CHATELIER.^ By Alfred J. Lotka. Received May 20, 1921. Presented by Raymond Pearl. The derivation of the principle of Le Chatelier from the laws of thermodynamics is familiar. We may approach a converse problem. What, in the broadest terms, are the conditions which a system must satisfy in order that the principle shall apply to it? The interest of this problem arises from the fact that we have reason to suspect these conditions may prove broader than the domain within which the laws of thermodynamics are conveniently applicable.^ We may therefore expect that a satis- factory solution of the converse problem may enable us to make rigorous application of the principle to systems to which, from lack of sufficient data it may be impossible, or from other causes it may be inconvenient to apply thermodynamic methods. Consider a system whose state is defined in terms of a variable x and a paramenter G. The system is one of that class, the history of which follows a law I = /(^^ G) (1) (For example, it may consist of a mixture of {A\— 2x) mols H2O vapor, (yl2+ 2a-) mols of hydrogen, and (^3+ x) mols of oxygen at 2000 deg. C. in a rigid enclosure of volume G; A\, A-i, A3 being con- stants, namel}^ initial masses). It is understood that other para- meters besides G may enter into the function /, but it is unnecessary to set them forth explicitly, since in the reflections which follow only 1 Papers from the Department of Biometry and Vital Statistics, School of Hygiene and Public Health, Johns Hopkins University, No. 37. 2 See Ehrenfest, Zeitschr. fiir phys. Chem. 1911, vol. 77, pp. 227, 244; Wolchonsky, Jl. Russ. Phys. Chem. Soc, 1912, vol. 44, pp. 305, 310; Chwol- son, Lehrbuch der Physik, 1909, vol. 3, p. 547; Bancroft, Jl. Am. Chem. Soc, 1911, p. 92; Fournier d'Albe, Contemporary Chemistry, 1911, p. 38; Lowy, Kosmos, 1911, p. 331; Le Dantec, La Stabilite de la Vie, 1910, p. 25; L. Fredericq, Arch, de Zool. Exp. et Gen., ser. 2, vol. 3, 1885, p. XXV; Spencer, First Principles, chapter 22, section 173, Burt's Edition, p. 433. For further historical and bibliographic notes see Duhem, Traite d'Energetique, 1911, vol. 1, pp. 523, 524. dx^"-' + I-- 5Xy 5G ' = - df /df ~ do/ dx 22 LOTKA. changes in x and in one parameter G at a time will be considered, the other paramenters being constant. According to (1) a stationary state (which need not be a true equilibrium in the thermodynamic sense) is defined by dx 0 = ^ = f{xr, G) ' (2) where Xi denotes the equilibrium value of x. If the parameter G is altered by a small increment 8G, the corre- sponding increment 8xi in the equilibrium value xi of x is, in view of (2), given by (3) (4) 1. Stable State. If the stationary state defined by (2) is stable, we must have in the neighborhood of that state ^ dx We can then distinguish two cases: fif a.) 7, > 0. This means that the parameter G is one whose in- crease accelerates the transformation the progress of which is meas- dxi ured by x. In this case it follows immediately from (4) that tt; > 0. oh In other words, if the s^\stem is stable in the stationary state defined by (2), then increasing a parameter which accelerates the transforma- tion will shift the position of the stationary state in the direction of increased transformation. From this alone, however, it does not necessarily follow that the new stationary state will actually become df 3 Condition (5) states that the velocity f = — 5x i.s alwavs opposite in dx sign to the (small) displacement 5x from equilibrium. This is evidently necessary for stability of equilibrium. VALIDITY OF THE PRINCIPLE OF LE CHATELIER.' 23 established. But, starting from the stationary state, at which dx — = f = 0, increase in G leads to a positive value of /. That is to dt say, a change actually takes place with a velocity directed towards the new stationary state, i,e. increased oc. b.) — :; < 0; i.e. increase in the paramenter G retards the trans- dG formation. Here it follows by similar reasoning that increase in G shifts the position of the stationary state towards diminished trans- formation. Furthermore, in this case the increment 56' initiates a retrograde change, i.e. a change toward the new stationary state. In both cases, (La) and (l.b), therefore, a change 8G in the para- meter G is followed by a transformation 8xi towards the new sta- tionary state, in the direction of the influence of the parameter G upon tl^ie velocity of transformation. 5, Unstable State. df Consider now the case in which — > 0. The stationary state de- dx fined by (2) is then unstable. A train of reasoning precisely analogous to that set forth above leads, in this case, to the conclusions: (1) A change 8G in the parameter G determines a shift of the stationary state in the direction opposed to the influence of the para- meter G upon the transformation. (2) The system, disturbed from existing stationary state by a change 56', moves, not towards, but away from the new stationary position. Application to Influence of Initial Masses. Consider a transforma- tion Sl+ ^2+ ...+ SrZ S'l-\- S'o+ . . . + S\ (6) Let ^1, ^2, . .^r be the masses (expressed in mols) at time t of the components Si, S2 . .Sr', similarly let ^'1, ^'2 -^'s be the masses of S 1, S 2- ■ ■ S s Let X measure the progress of the transformation from left to right, and let pi x be the amount . (in mols) of S, transformed from time t = 0 to time t = t. Let Ai be the initial value of the mass (in mols) of some component 24 LOTKA. Si which disappears in the transformation when x increases, and let A'j be the initial value of some component 0 for every component which dis- df appears in the transformation, and if z~Tr < 0 for every component oA ] which appears, then, in view of (9), the same is true of -rr and ttt . But dii = -pidx (11) d^'i = + p'idx (12) where p», p', are positive numbers, and dx~ ^ d^i dx ^ ^ a^y dx ^^^^ = - i:§Pi+ zSv'i (14) which is necessarily a negative quantity if ,-^>0, ^-i<0 (15) 3. It should be noted that the argument by which our conclusions Lave been drawn depends on the existence of equations of constraint, relations such as (8), connecting the ^'s and the A's. In the absence 26 LOTKA. of such constraints we are in no wise assured that the principle holds.* This must be clearly borne in mind in seeking to apply the Le Chate- lier principle, for example, to biological systems. Thus, for instance, the malaria equilibrium under the conditions contemplated by Sir Ronald Ross,^ is independent of the initial amount of malaria in the system (provided only this is not zero). This state of affairs arises out of the fact that there is no equation of constraint of type (8), in this case, connecting the initial amount of malaria with its status at any subsequent epoch. Case of more than one variable. A somewhat more complicated case arises if the system under consideration is susceptible of several con- current transformations, so that its state at any instant requires for its definition not one variable x, but a number of such variables. It will suffice if we consider here the ca.se for two variables x, y, as, for example, the case of a pair of consecutive reversible reactions In this case we have dx n't ~ /i (•'•- y,o) dy dt h (x, y,G) and equilibrium is defined by /i = h= -- 0 AZ ^^Z (' (16) (18) (19) (20) Differentiating, in a manner analogous to that followed in the case of a single variable .r, we have ^U a/i dfr - P)U du a/o ~ 6a-, + t^ 6?/i + Jf, 6 6' = 0 (22) dx dy dU 4 For there is then no necessary relation between $ and A, so that the r)f Hf derivative -^ is no longer equal to ^, but is indeterminate or meaningless. 5 "The Prevention of Malaria," Second English Edition, John Murray, London, 1911, p. 679; Lotka, Nature, Feb. 1912, p. 497. VALIDITY OF THE PRINCIPLE OF LE CHATELIER. 27 5.ri 5//1 a system of linear equations, which we solve for — , ^y; and obtain dh 5/1 dG dy dh a/2 8xi dG dy 8G 5/1 dfi dx dy dJ2 dx dy (23) 8yi and a similar expression for 777 . Condition of Stability. A general solution of (18), (19) can be written ^ in the form of exponential series x= Poi- Pie"'' + Poj''^' + Pne^^'' + where Xi X2 are the roots of (24) (25) A(X) ,6x dx - X ^df, \dy dJ^ dy -X 0 (26) The condition for stability "^ of the equilibrium is that the real parts of all the roots X are negative. This in turn demands that the abso- lute term A(0) be positive. But this absolute term is, evidently, A(0) dfi dfi dx dy dh df2 dx dy (27) 6 A. J. Lotka, Proc. Am. Ac, 1920, p. 139. 7 Idem, loc. cit., p. 144; Hurwitz, Math. Ann., 1875, vol. 46, p. 521 ; Blondel, Jl. de Physique, 1919, pp. 117, 153. 28 LOTKA. SO that we must have, for stability, 5/1 5/1 d.r dy dfo a/2 dx dy >0 (28) In consequence, given stability of equilibrium, the sign of dxi dG will be the same as that of the numerator in (23), i.e., that of the expression " dG dy dG dy (29) Example. Consecutive Reactions. By the way of example we may apply these results to the case of a pair of consecutive reversible reactions. Si+ S2+ . . . + SC S'i+S'2+ . . . + S'C «"!+ ^"2+ . . . + S"t (30) Let X denote the progress of the first reaction from left to right (so that, for example, a quantity, -piX of the substance S» has been trans- formed at time t); and let y similarly denote the progress of the second reaction, from left to right. Let us consider the effect upon x\, the equilibrium value of x, of an increment bA" k in the initial amount of substance S" k appearing as product of the second reaction. We have, according to (23) dh a/i dA"u dy dh a/2 bxx dA'\ dy bA'\ dfi a/i dx dy a/2 a/2 dx dy (31) We shall assume stability, so that the denominator is positive. In the numerator, evidently ^ 8 If we exclude any possible catalytic influence. VALIDITY OF THE PRINCIPLE OF LE CHATELIER. 29 so that this numerator reduces to = 0 . (32) Stability demands dA"k dy f > 0 .(34) On the other hand the principle of Le Chatelier would make This, by (31), in view of (32), (33), (34), will be true or not according as S^ > 0 (36) Hence the principle of Le Chatelier holds good or not, as applied to the effect of A"k upon xi, according as af-J" (3^) From this point on the discussion would follow essentially similar lines as in the case of a single dependent variable; it is therefore un- necessary to carry this out in further detail. Influence of External Factors. We have hitherto tacitly assumed that (1), or (18), (19) are the only conditions for equilibrium, or, that, if there are any other conditions to be satisfied, these are in some way automatically taken care of. In point of fact, in general, in addition to a condition of the form Jt = ^^'' ^^ ^^^ there will be further conditions of the form H= H, (38) where H is a, parameter entering into the function /, while //« is a parameter defining certain "external conditions." For example, H may be the pressure exerted by a gaseous mixture against an enclosure, and He may be the external pressure applied to a movable piston 30 LOTKA. forming part of that enclosure. Here it is not enough, for complete equilibrium, that (1) be satisfied, but (38) also must hold. Furthermore, the conditions (1) and (38) define equilibrium, but are insufficient to determine its stability, since they give us no in- formation regarding the behavior of the system when H + He, i.e. when not in equilibrium with the external parameter H e- In order to settle this point we must have some further data. We are here interested in systems in which such additional data are furnished in the following manner: In the case of these systems it is found that, in relation to the parameter G a certain parameter H ha\nng certain peculiar properties, can be defined by a relation. ip (^1, ^2, G,H) ^ constant (39) or its equivalent -^ {x, Ai, A^,. .G,H) = constant (40) The peculiar propert\' of G referred to above is as follows dG>_ > —-. 0 according as // — //« ^=0 (41) dt < ^ < It will perhaps be well, before proceeding any farther, to illustrate this by a concrete example. Consider the system 2 //2O ^ 2 H2-\- O2 (42) If ^1 is the mass of II 2O expressed in mols, ^'1 the mass of Ho and ^'2 the mass of O2 similarly expressed; if I' is the volume (parameter 6-') and if P is the pressure (parameter H) exerted upon the enclosure, then the equation (39) here takes the form py = (^1+ ^1+ r^) Re (43) where 6 is the absolute temperature and R the general gas constant. Or, if ^1, A'l, A'2 are the initial masses of H^O, Ih and O2 respectively, (expressed in mols), and x measures the progress of the reaction, as, for example, by the number of O2 mols formed, then evidently (44) (45) (46) .^1 = Ar — 2x l'l= A' 1+ 2x ^2 = A' 2+ X VALIDITY OF THE -^PRINCIPLE OF LE CHATELIER. 31 SO that (40) takes the form PV = { Ui- 2x) + (.l'i+ 2x) + {A'o-\- x) Rd ..^. = \{x + A,+ A\+A'^^Re ^^^' In this case it is quite evident that the parameters P, V (correspond- ing to //, G of the general case) have the property defiiifd by (41), which here appears as the characteristic property of the intensity factor and the capacity factor of an energy. But for our present purposes we are not concerned with the question whetjier or not the parameters G, H defined for a given system are or are not factors of an energy. We must be prepared to deal with cases where this is either uncertain or actually known not to be true. All we need to know, for our purpose, is that the parameters G, H have the property defined by (41). An example may serve to illustrate the fact that this property may be shared by physical quantities not obviously related to energy. Among the parameters on which the rate of increase of a human population depends is the area a occupied by them, since this deter- mines the population density, which in turn influences the death rate in well-known mann^r,^ and. presumably, in some degree the birth rate also. Now there is an obvious relation between population density and ground rent. Regulation is effected about as follows: There is a certain demand for space, a desire for expansion, which may be measured by the rent // per unit area that the individual is willing to pay. On the other hand there is a certain market price He which must be paid to obtain accommodatioji. Now \i H > II e, i.e. if, on an average, an individual is willing to pay more than the market price, the population will spread over a greater area by renting more ground. If, on the other hand // < II e the individual is not willing to pay the market price, he will retrench, he will move from a six room apart- ment to a five room apartment say, and the area occupied by the population will contract. The parameter He functions, in fact, much like a "surface pressure," tending to compress the population into a smaller area. The most striking exhibition of this " surface-pressure " is seen in a great metropolis such as New York, where the population, a naturally two-dimensional structure spread like a film over the earth's surface, has been thrown into great creases towering 700 feet and more, 50 layers deep, above the street level. 9 See, for example, Newsholme, Vital Statistics, 1899, p. 154. 32 LOTKA. It will be seen that in this case the internal parameter H and the corresponding external parameter He so determine changes in the area a that da> , > — -: 0 according as H — He~ 0 (48) that is to say, the parameters //, a and He are related to each other and determine the course of events in a manner analogous to the intensity factor, the capacity factor of an energy, and the "applied force." But it is quite unnecessary to suppose that H and a actually are such factors of an energy in the example cited (population-spread) ; on the contrary, the writer is opposed to this view, which he has taken occasion elsewhere to discuss. ^° For our purposes it is quite immaterial whether P and a are factors of an energy. All we need know is that they enter into the condition (41) as there set forth. Condition for Stability toward External Factor. Consider a system for which the condition for equilibrium with the environment is given by H = He (p (G, H) = constant dG > ,. „ rr > —r = 0 aceordmg as H — He = 0 at < < (49) Let

Hi (52) > He (53) 10 "Economic Conversion Factors of Energy," to appear in a forthcoming issue of Proc. Nat. Ac. VALIDITY OF THE PRINCIPLE OF LE CHATELIER. 33 Th en dG dt >0 (54) Hence the point moves along the curve in the direction Ai A2, i.e. still farther away from equilibrium. On the other hand, the same reasoning applied to a curve sloping downward from left to right shows that the system after displacement returns to its equilibrium position. So, for example, the curves representing the relation between pres- G Figure 1. sure and volume of a gas necessarily slope downward from left to right; the same is true of the demand and supply curves of economics. If it were true, as sometimes stated, that the more a man has, the more he wants, economic equilibrium would be an unstable condition. External Stability and the Principle of Le Chatelier. Consider now a system which obeys the condition ^ {x, G, Hy= 0 ^- = 0 according as H — He^O at < < (55) (56) 34 LOTK.^^. Let the system be stable towards He both when x is held constant and also when a- is at the equilibrium value a-i defined by J^=f (^-1, 0) = 0 (57) This means that all the curves " of constant composition " if {x, G, H) =0 j (58) X = constant j and also the curve " of equilibrium composition " cp (xi, G, II) = 0 (59) slope from left to right downwards. Now consider two neighboring curves of type (58) (curves of con- stant composition), which we will suppose solved for H and write 7/„=^a(6'. a-„) (60) H,==-^b{G,x,) (61) SUppose we start with the system in the state represented by the point Q, in internal equilil)riuni and also in equilibrium with an ex- ternal parameter He (see Fig. 2). Let x be changed at constant G, so as to increase H according to (55) until X = Xb, so that the representative point strikes the second curve of constant composition at R. Since at the start of this operation // = //^= //^ (62) and at the end H = Hbi > H, \ f63) therefore the system is not in equilibrium with the external pressure He in the state represented by the point R, but equilibrium (for X = Xb) occurs at some other point T which must lie to the right of R along the curve of constant composition RT, since, whenever // >He G increases, in accordance with (56). Furthermore, drawing a horizontal QS, T must lie below S, since the line of equilibrium composition QT must slope from right to left downwards. VALIDITY OF THE PRINCIPLE OF LE CHATELIER. 35 It is clear therefore that lines of constant composition are steeper than lines of equilibrium composition. It follows at once that if G be increased while the system is kept in equilibrium, so that the representative point travels along QT, then the change in x from Xato Xb is that which at constant G increases H, or at constant // increases G. But this is the principle of Le Chatelier. This principle therefore Jiolds whenever the conditions (55) (56) are satisfied, and the system H G Figure 2. is stable towards Hg both when x — constant and when x = xi (i.e. when x has its equilibrium value). Similarly, it can be shown that if the conditions (55), (56) are re- placed b^- . - ,

_ -77 — 0 according a,s H — He — 0 dt > ^ < 36 LOTKA. while at the same time the system is stable towards H both when a- = constant and also when x = xi, then the Le Chatelier principle holds. In this case the curves

S hkuh;ma\. soom lo luo ihat tluMV is a j;ivat doal of vsigiiifioancv in iho '" oooilioiont of spooitio irsisiauiv." Howovor, it is of intorost lo noio in ilic tahlc that tho changvs of diuieiisioiis of maiiijaniti ami tluM'lo aiv so iaruo v'oiu[>aivil witli tho tonsimi oooHioiom of obsorxod rosistamv tliat tho toiision v'oothoiont of spooitic rosistaiioo is iiogativo, wlioroas tho tonsioti iwlUoioiu of obsorvo«i rosistanoo is positivo. For tho othor inoials tho ciM'rtvtiotv for ohaiijjo of figiuv is not larj^o oiiongh to ohango tho sign of tho a>othoiont of obsorvoii n\sistaiioo. It is ill tho first phioo to ho romarkotl front tho tahlo that of tho sovon substattcvs whioh aro ahtiornial wiih rospoot to tho siirn of tho prossttro ivotlioiont. only two, bismuth and strontimn. aiv abnormal with n\spoor to tho sign of tho tonsion cH>otticiont. This would soom to indioaio somo ossontial dilVoronoo botweon tho oondtiotion moohanism of thoso two stibstamvs and that of tho othoi-s. Lot ns disonss what this ditVotxMtcv may bo in tho light of tho thtvry of motallio oondnotiott whioh 1 ha\o ptvvionsly do\olopod. 1 havo thought of ciwduotion as duo to a froo path mtx'hanism; tho olassioal thtx^ry was a frtv path ihoory. Tho ditVortnicos oomparod with .tho olassioal tlu\M"y aro those. In tho first place, tlie free paths aro thotight of as loitg, booanso tho frtv olootivns an^ fow in ntimbor. In normal motals. tho paths of tho oUvtn.>ns aro to bo thought of as tlnxnigh tho snbstancv of tho atoms thomsolvos. The path may bo terminattHi w hon tho eleotUMi makes the jnmp frtun one atom to the next. Tho ohanct* of termination on making the jump will depend both on the amplitude of atomic vibration and tho distantx^ apart of tho atoms. Now if tho distamv apart of the atoms varies little com- patwl w ith tho ohangi^s of amplitude, the variation of ft\x^ path may be calcnlatoii in terms of the variation of amplitude only. The changes of amplitude, nogkvting the ort'eots dne to changes of dimensions, may be calcnlatoii for changes of prt^s^nre and tomperatnre, and so the ehangi^ of path, and hoiuv tho changt^s of rosistamx^ may also be cal- cnlaiixl. It is in ihrc»wing the entire bunion of tho variations on the free p o ^ o z o o < 3900 3600 3700 4 5 6 7 8 Pressure. Kg./Cm.'X 10^' Iron 10 li 12 Figure 6. Iron. Thermal conductivity on an arbitrary scale against pressure in thousands of kg/cm-. Results obtained with a longitudinal flow specimen. The points he on several hnes of the same slope; the reason for this is explained in the text. with the three good samples, after applying all corrections, were —0.6, —0.2, and —0.2% respectively. The discarded data were not inconsistent with these values. We take as the mean —0.3%. The pressure coefficient of thermal conductivit}" given by the above results is — O.OeS. I found for the pressure coefficient of electrical conductivity between 0 and 12000 kg. +0.0o229. Iron is not one of the metals measured by Lussana, so there are no previous values for comparison. EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 109 Copper. Two specimens were made for the radial flow method, and four for the longitudinal. As in the case of iron, the thermo-couples and heating element of the radial flow specimens were placed in copper tubes sweated into place, and the results were not at all satis- factory. The points were not regular, and the irregularities repeated, showing some real effect. Furthermore, the two radial flow specimens were made from contiguous lengths from the same piece of commercial drawn rod, and the irregularities were much the same in character for each specimen, showing a real effect of inhomogeneities in the metal. The four longitudinal specimens were made from electrolytic copper wdiich I obtained a number of years ago from the Bureau of Standards. 3 4 5 6 7 8 Pressure. Kg. / Cm/ X 10' Copper Figure 7. Copper. Thermal conductivitj^ on an arbitrary scale against pressure in thousands of kg/cm-. Results obtained with a longitudinal flow specimen. Their analysis is as follows : Cu 99.995 per cent, trace S, no x\g, CU2O, As, or Sb. It is to be noticed that the purity is unusually high. Measurements were made on some of these samples in the annealed condition, and others not; there seemed to be no difference in the results. These four samples gave fairly good results, the points lying on discrete lines, as usual. The least scattering of these is reproduced in Figure 7. The effect is seen to be fairly large, and negative, the mean effects shown by the four samples were —9.7, —8.2, —8.3, and — 7.5% change respectively under 12000 kg/cm^. The mean is —8.4%, but we will take as the best result —9.0% instead, because 110 BRIDGMAN. the best of the specimens gave the higher results. Because of the high conductivity of copper, the correction for the transmitting medium is small, being only 1.2%. The pressure coefficient of thermal conductivity deduced from the data above is — 0.0o75. No departure from linearity could be de- tected. I have found for the effect of pressure on electrical con- ductivity over this range the mean value +O.O5I8I. Lussana's results for copper run only to 1000 kg. He found the effect to be linear, and the conductivity to increase under pressure, the opposite sign from my results. His coefficient is +O.O5IO. The relation be- tween pressure and electrical resistance he also found to be linear, and the coefficient to be O.O5212. Silver. Only the longitudinal flow method was used on this metal. The material was obtained from Baker, and was said to be of high purity, but I have no analysis. Three different samples were used. The two runs made on the first two samples apparently indicated an increase of conductivity- under pressure. The reason for this has been discussed in detail previously; not enough readings were taken, and there was a tendency for the points to shift from a lower lying to a higher lying curve with increase of pressure, thus simulating a false effect. IMore readings were made on the third sample, and an effort made to control the position of the points on one or another of the possible lines. The results with this sample are reproduced in Figure 8. The tendency of the points to lie on one or another of three dis- tinct lines is obvious, as also the fact that the effect is negative, the conductivity decreasing with increasing pressure, instead of increasing, as was indicated by the results first obtained. The two first samples were now set up again, and the measurements repeated, with the precautions which had been gained from the intervening experience. The results shown by these two samples were now also unmistakably negative, and of nearly the same numerical value as shown by the third sample. The numerical magnitude of the total decrease of conductivity under 12000 kg/cm^ shown by the three samples was 4.1, 4.4, and 4.6% respectively. The correction for the pressure effect on the transmitting medium was 1%. The pressure coefficient of thermal conductivity given by the above data is —O.O^Sl. The average pressure coefficient of electrical conductivity between 0 and 12000 kg. I have found to be +0.05334. This metal was not investigated by Lussana, so there are no previous results for comparison as to the thermal conductivity. Nickel. Both methods were used for this metal, and material from EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. Ill two different sources. I am indebted to the International Nickel Co. for a piece of | inch round rod of high commercial purity (99%). From this two samples were made for the radial flow method, large cylinders without the sheath. The thermo-couples and heating ele- ment were placed in fine copper tubes sweated into place. The re- sults obtained with these samples were very irregular, and it was evident that the thermal contact between the copper and the nickel was not sufficiently good. These measurements did little more than establish a strong probability that the effect was negative. After the radial flow measurements had been made, two small pieces were 0 12 3 4 5 6 7 8 9 Pressure, Kg. / Cm.' X 10' Silver « FiGTTRE 8. Silver. Thermal conductivity on an arbitrary scale against pressure in thousands of kg/cm-. Results obtained with a longitudinal flow specimen. The points lie on several lines of the same slope; the reason for this is explained in the text. cut from one of the cylinders for longitudinal flow specimens, and a few readings were made with them. This was before the explanation of the scattering of the points by this method had been found. The results were very scattering, and repetition would have been necessary to make sure of even the sign of the effect. After completing the measurements on commercial nickel I was fortunate enough to obtain through the kindness of Mr. I. B. Smith of the Research Laboratory of the Leeds and Northrup Co. several samples of exceedingly pure nickel. I have no analysis of the nickel, but its high purity is vouched. for by the unusually high value of the temperature coefficient of electrical resistance, which between 0° and 112 BRIDGMAJSr. 100° was 0.00634, against 0.0049 for commercial nickel over the same range. Two longitudinal flow samples were made from this pure material. The first of these samples gave points lying on tliree different lines separated by the usual 5%. The slope corresponded to a decrease of conductivity of 13.5% for 12000 kg. In setting up the second sample I made the attempt to prevent motion of the thermo-couple wires in the holes with a piece of 0.002 inch wire laid beside them, as has been >- > a z o o < i llJ X 1900 1850 1800 1750 1700^ 4 3 6 7 8 Pressure. Kg./Cm.'X 10' 10 II 12 Figure 9. Nickel. Thermal conductivity in arbitrary units against pressure in thousands of kg/cm^. Results obtained with a longitudinal flow specimen. The points lie on several lines of the same slope; the reason for this is explained in the text. mentioned in the description of the method. This was the first attempt, and as often happens, succeeded better than subsequent attempts. For some reason I was fortunate enough to get the wire into place without introducing strains into the thermo-couples, and the results showed a gratifying regularity. The results of the final run with this second specimen are shown in Figure 9. It is seen that the readings still lie on three different lines, but these lines are now E^ECT OF PRESSURE ON CONDUCTIVITY OF METALS. 113 separated by much less than 5%, as is to be expected. A partial run with this same specimen, which had to be discontinued because of leak and also because of short circuit in the three-terminal plug, gave exactly the same slope for those readings which could be obtained as the final run. The change shown by this second sample was a decrease of conductivity of 14.5% for 12000 kg., agreeing fairly well with the first sample. Since the second sample gave somewhat more regular results, it is given greater weight in the mean, which I take as 14.1%. The total correction for the effect of pressure on the trans- mitting medium was 5% of the total conductivity, amounting to about 33% of the observed change under 12000 kg. The results found above give for the pressure coefficient of thermal conductivity — O.O4I2. There are no previous results for comparison. An incidental result obtained from the measurements with com- mercial and pure nickel was a comparison of the absolute thermal conductivities. The longitudinal flow method is not well adapted to give the absolute conductivity because of the uncertainty in the cor- rections for loss through the leads, etc. (the absolute conductivities directly calculated average about 5% higher than the values of Jaeger and Diesselhorst), but the comparative values of absolute conductivity of different materials should be nearly correct. The thermal conduc- tivity of the two samples of pure nickel was found to be 37% higher than that of the two samples of commercial nickel. Considerable confidence may be put in these values, as the individual readings were very consistent; the two samples of pure nickel gave results differing by less than 0.5%, and the results on the two samples of commercial nickel were identical to three significant figures. Platinum. Measurements were made on two different samples by the longitudinal flow method. The material was obtained from Baker, the purest which they could supply. I have no analysis, but have the statement of Baker that the impurity is guaranteed to be less than 0.1% and is probably not greater than 0.01%. The measurements on platinum were scattered on three lines of a maximum separation of about 5%, as is usual with this method. The two specimens gave identical results, a decrease of conductivity of 1.9% for a pressure change of 12000 kg/cm^. The observed effect was positive, but the correction for the effect of the transmitting medium is so large, 5.2%, as to alter the sign of the result. The pressure coefficient of thermal conductivity as given by the above measurements is — O.O5I6. The pressure coefficient of electrical conductivity at 30° between 0° and 12000 kg. is +O.O5I86. 114 BRIDGMAN. Bismuth. An attempt was made to obtain measurements on this metal by the radial flow method, but without success. It did not prove possible to get sufficiently good thermal contact with the fine copper tubing, and I have already mentioned that the attempt to use silver tubing failed because of the extremely rapid alloying action between silver and bismuth. Measurements were finally made on three different specimens by the longitudinal flow method. A great deal of time was spent in the endeavor to obtain pure material. In my previous work on the effect of pressure on electrical resistance I had purified the metal by electrolysis. I now endeavored to repeat this, but without success; I could not make the bismuth form a coherent deposit. The procedure of the previous work was exactly repeated as far as I could tell. Previously the hydrosilico- fluoric acid had been obtained from a German source; this was no longer available and acid from the J. T. Baker Chemical Co. was used instead. The acid was of high purity as indicated by the analysis on the label, but there is a possibility that some impurity not covered by the analysis might have been responsible. I then obtained some bismuth from the U. S. S. JNIetals Refining Co. I have to thank them for supplying me with six pounds of the metal, in two lots. Their product is prepared electrolytically, in distinction from the ordinary commercial product, and is guaranteed b\' them to have less than 0.1% impurity. Ordinary commercial bismuth has about 3% im- purity. My test for purity has been the temperature coefficient of electrical resistance. This electrolytic bismuth showed a very low coefficient, only 0.0022 between 0° and 30°. Ordinary commercial bismuth is higher. Professor F. A. Saunders was kind enough to make a spectroscopic analysis; he found a very strong silver line, which seemed to indicate a rather considerable impurity. I con- sulted the U. S. S. Metals Refining Co. again, and they were so kind as to send me a second sample, which they had submitted to chemical analysis, and found to contain only silver in detectible quantity, and this was less than 0.06%. But the temperature coefficient of this new sample was again very low. I attempted a purification by slow crystallization from the melt, with the result of bringing the coefficient up to only 0.0025. Ordinary commercial material, purified in the same manner, showed a coefficient of 0.0034. Professor Saunders was again kind enough to make a spectroscopic analysis and found again the strong silver line, which seemed to him could only arise from a rather large amount of impurity. He found traces of Cu and Pb, and no traces of Sn, Cd, Zn, Li, As, or Sb. Professor G. P. Baxter EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 115 was now so kind as to make a quantitative determination of the silver, and found 0.03%, confirming the conservative estimate of the U. S. S. Metals Refining Co. I now succeeded in finding a small residue of my original electrolytic bismuth, and Professor Saunders made a spectroscopic analysis of this. He could find only traces of Cu and Pb, the Cu being stronger than in the commercial electrolytic bismuth, and some silver, evidently considerably less than in the commercial. The conclusion seems forced that a quantity of silver as small as 0.03% can depress the temperature coefficient to half the normal value, thus exerting an effect very much greater than such impurities as Pb and Sn, which are present in ordinary commercial bismuth. That difficulty would be expected in removing the silver by recrystalli- zation is evident on an inspection of the mixed crystals diagram for these two metals. This would also be indicated by the energetic alloying of silver and bismuth, which made impossible the preparation of the radial flow specimens. Under the circumstances it seemed that the best thing to do was to use the commercial electrolytic bismuth, with its known analysis of 0.03% of silver, in the expectation that the effect of this small impurity is abnormally high on the temperature coefficient of resistance. I had previously found that the effect of impurity on the pressure coefficient of resistance is much less than on the temperature coefficient of resistance. The samples were made from | inch wire which had been formed by hot extrusion in the regular way. One advantage of forming the specimen by extrusion is that the crystalline structure is very much finer than when the specimen is cast, and so the results are much more likely to give the average for all the directions of a single crystal. The thermo-couple holes were drilled in these specimens in the regular way, but a modification was necessary in mounting the heating element. Previously the heating element was mounted in a copper capsule, which was cemented into a hole drilled in the end of the specimen. This was no longer possible, because the capsule was so large that it was not possible to drill a hole to receive it without breaking out the walls in so brittle a material as bismuth. The heating element was accordingly placed in a smaller hole drilled directly in the end of the specimen. This has the disadvantage that the terminal conditions of temperature are not so accurately defined as with the other metals, and the motion of the heating element in its receptacle may produce other irregularities. This was indeed the fact; the points were more scattered than with the other metals, and 116 BRIDGM.^N. the width of the band of scattering was greater than the 5% usually found. The magnitude of the scattering varied with the specimen, as it might be expected to. In setting up the first sample, the ground of the heating element was made to the sample itself. It has been previously exphiined that this introduced an effect due to the Peltier heat, which is unusuallv large for this metal. The mean of readings with two directions of the heating current should eliminate this effect. With the other two samples an independent ground was used, and the eft'ect disappeared. It is to be expected that if there is any error due to heat leak along the thermo-couple wires that it will be especially large for this metal, whose own thermal conductivity is so small. In order to test this point, duplicate runs were made on the third sample, first with the ordinary copper-constantan couple, and then with a couple of " therlo" constantan. Very nearly the same results were found, showing the adequacy of the precautions taken to prevent leak along the couple wires. The third sample gave the most regular results; probably some accidental twist or bend in the wires made them less likely to be displaced under pressure than those of the other samples. The results with this sample are reproduced in Figure 10. The results found with the different samples are as follows:^!, -38.8% for 12000 kg.; ^2, -38.8%; #3 (copper-constantan couple) —37.3%, and ^'3, (therlo-constantan couple) — 35.5%. Mean —37.8 %. Because of the low conductivity of bismuth the correction for the transmitting medium is large, amounting to 13.8%, and is in the direction to make the true effect more negative than the observed effect. The pressure coefficient of thermal conductivity to be deduced from the above measurements is — O.O43I. It is to be noticed that this is negative, and also that it is abnormally high. The abnormal sign agrees with the pressure effect on electrical conductivity, which also decreases under pressure. I found at 30° the average pressure coefficient of electrical resistance up to 12000 kg. to be — O.O4212. Bismuth was not among the metals measured by Lussana, so there are no previous results for comparison. Antimony. IVIeasurements were made on four samples, all by the longitudinal flow method. The material was obtained from the J. T. Baker Chemical Co. It was supposed to be especially pure, although I have no analysis. Antimony from the same source was formerly used by the Bureau of Standards to give a fixed melting point, but EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 117 their experience was that although the chemical analysis might show very little impurity, there was nevertheless present in all antimony from American sources some slight impurity which was sufficient to displace the melting point by several degrees. Presumably my anti- mony suffered from the same impurity. Two of my samples were made from cast antimony and two from extruded metal. The metal was cast by pouring it into a groove in a massive iron block to a thickness of a trifle over | inch. The chilling was hence very rapid, and the crystalline structure very fine. From 1500 0 10 12 3 4 5 6 7 8 Pressure, Kg. / Cm.' X 10 Bismuth Figure 10. Bismuth. Thermal conductivity in arbitrary units against pressure in thousands of kg/cm^. Results obtained with a longitudinal flow specimen. this casting two pieces were machined for the longitudinal flow method. The metal is so brittle that it was not possible to cut it with the tool in the ordinary way, but the machining had to be by grinding. Even then extreme caution was necessary, and there were many fail- ures before success was attained. The wire for the other two speci- mens was extruded at a red heat through a high speed steel die of the required dimensions. Some little practice was needed before the proper manipulation was found. It is possible to extrude antimony at a considerably lower than a red heat, but with a wire as large as | 118 BRIDGMAN. inch the product is Hkely to be very brittle, or break spontaneously into small pieces. If the temperature is raised very close to the melt- ing point, however, it is possible to get by extrusion a uniform straight wire with no apparent flaws, and not as brittle as the casting. The thermo-couple holes were drilled in the four pieces in the regular way. The heating elements, as in the case of bismuth, were mounted directly in small holes drilled in the ends, it not being feasible to drill so large a hole as the use of the copper capsule would have demanded. This manner of attaching the heating element was responsible for the greater irregularity of the points, as also in the case of bismuth. In one case the scattering was such and the accidental 0 I 2 3 4 3 0 7 ■-) Pressure. Kg./Cm.'X 10' Antimony Figure 11. Antimony- Thermal conductivity in arbitrary units against pressure in thousands of kg/cm-. Results obtained with a longitudinal flow specimen. distril)ution such that a positive sign for the effect might have been suspected. The readings obtained with the first of the cast specimens were the most regular; these are reproduced in Figure 11. The thermo-couple used with the second of the extruded specimens was therlo-constantan, instead of copper-constantan. The readings with this were essentially the same as with the others, thus again showing freedom from heat leak along the wires of the couple. The thermal conducti\dty decreases with rising pressure; the two cast specimens gave respectively —23.9 and —26.3%, and the two extruded specimens —24.8 and —23.9%. The mean of all four is EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 119 —24.7%. It is to be noticed that within the limits of error no differ- ence is to be detected between the cast and the extruded specimens. The pressure correction for the transmitting medium was 15.3% on the total conductivity; this means that the final corrected result was three times as large as the observed pressure effect. The above results give for the pressure coefficient of thermal con- ductivity — O.O42I, larger than for any other metal except bismuth. I have previously found that the electrical conductivity of antimony also decreases with rising pressure, and at 30° the average coefficient to 12000 kg. is -O.O4IO8. Lussana has also measured the effect of pressure on the electrical and thermal conductivity of antimony, and his results are in precise disagreement with mine. He finds that the electrical conductivity increases under pressure, as it does for normal metals. At 25° his coefficient, presumably to 3000 atmospheres, is O.O5874. There must be something vitally wrong here; measurement of electrical resistance under pressure should offer none of the difficulties of thermal conduc- tivity, and there should be no reason for a disagreement as to sign between different observers. The relation between pressure and thermal conductivity Lussana finds to be distinctly not linear. The initial change is at a rate corresponding to a coefficient of O.O525I, and at 3000 the rate corresponds to a coefficient of only O.O5I64. So large a departure from linearity in a metal with as high a melting point as antimony is without precedent. It does not seem improbable that the sign that Lussana found for the coefficients of both electrical and thermal conductivities may be due to a closing of minute fissures between the crystalline grains under pressure, such as Borelius and Lindh found for bismuth.^ The departure of his thermal conductivity from linearity is in accord with this suggestion. Petroleum Ether. It has already been explained that it was neces- sary to determine the absolute conductivity and pressure coefficient of this substance in order to obtain the correction due to the effect of pressure on the transmitting medium in the longitudinal flow method. The method adopted for determining these constants for petroleum ether was a radial flow method, and demanded very little change in the apparatus already used for metals. The apparatus is shown in Figure 12. It consists of an inner cylinder of copper held concen- trically within an outer hollow cylinder, which in turn fits closely inside the pressure cylinder, and is maintained concentric with it by the same spring device that was used for the metals. The petroleum ether fills the annular space between the two copper cylinders. The 120 BRIDGMAN. axis of the central cylinder contains a linear source of heat, that is a wire carrying a current, precisely as for the metals. The difference of temperature between the outer surface of the inner cylinder and the inner surface of the outer cylinder is measured by thermo-couples. These were of the same construction as for the metals, and were mounted in fine copper tubes, which were sweated into small holes drilled lengthwise of the cylinders. Of course with this construction the couples could not be located exactly on the surface of either cylinder, but the thermal conductivity of the copper is so much higher than that of the petroleum ether that practically all the temperature drop takes place across the annular space of the liquid, and a rough computation shows that with the dimensions used any error from this cause is negligible. As a precaution against failure of perfect geometri- inn\unnnumn\vnHi«i»\\muuun»\H\nu»nunuu\u\u\»n IMwmwwuwwwwvwwwwwwwwwwwwuwwwwTOwnwmuwuwn Figure 12. Longitudinal section of the apparatus for measuring the thermal conductivity of petroleum ether. The liquid is showTi shaded between copper cylinders, with a central heating unit and two sets of thermal junctions bridging the liquid. cal centering of the inner cylinder in the outer one, three couples instead of one were used, spaced at even angular intervals around the cylinders, and these were connected in parallel, so that the reading obtained gave the mean of the temperature differences around the cylinder, and any geometrical imperfection is eliminated. The annu- lar space between the cylinders was only 1.3 mm. wide. This is so narrow as to remove any error from convection in the liquid, even at atmospheric pressure, and it has already been explained that such error vanishes at higher pressures because of the rapidly increasing viscosity. No effects were found in the measurements to suggest error from such a source. Because of the substantial equality of temperature throughout the copper cylinders, it is to be expected that errors from slight changes in position of the thermo-couples, which played so large a part in the measurements of the metals, would vanish. This is indeed the fact, and the measurements showed a high degree of regularity, much beyond that obtained for any metal. In making the readings, the entire interior of the apparatus was EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 121 filled with petroleum ether. The same method would serve for any other liquid which does not absorb impurity, or become conducting under pressure. Unfortunately most of the liquids whose other properties are best known, and which it would be most interesting to measure, such as the alcohols, are not of this kind. To determine the thermal conductivity of these under pressure it will be necessary to so modify the apparatus that the liquid can be insolated from the electri- cal leads. I 2 3 4 5 6 7 8 9 10 II 12 Pressure, Kg. /Cm.' X 10' Petroleum Ether Figure 13. Petroleum Ether. Thermal conductivity in arbitrary units against pressure in thousands of kg/cm-. Results were obtained with the apparatus of Figure 12. The observed results with petroleum ether are shown in Figure 13. The greater regularity of th'fe results as compared with the metals is manifest. The lowest pressure of these readings was 500 kg. The reason for not going to atmospheric pressure was not error from con- vection currents, but because at this pressure the heating effect would have been sufficient to vaporize the ether, and so introduce error. 122 BRIDGMAN. The effect of pressure is seen to be a large increase of conductivity, amounting at 12000 kg. to an increase of 2.22 fold. The increase is not linear with pressure, but there is a departure from linearity in the normal direction, in that the change becomes proportionally less at the higher pressures. The initial rate corresponds to an increase of conductivity of about 20% per thousand kg. So far as order of magnitude goes, this agrees with Lussana, who found the correction for his transmitting medium to be at the rate of 30% per thousand kg. He did not find a departure from linearity. Of course there is no reason to expect more than agreement as to order of magnitude, because his transmitting medium was a comparatively heavy oil, quite different in properties from mine. The measurements of the effect of pressure on the thermal conduc- tivity of liquid is a thing worth doing for its own sake, and I hope to get the chance to make measurements on a number of others. In fact I already have results for two alcohols and kerosene. Suffice it here to mention that there seems to be an intimate connection between the pressure effect on thermal conductivity and the pressure effect on the velocity' of propagation of sound. General Comment on Lussana's Results. The only previous measurements of the effect of pressure on thermal conductivity are those of Lussana. Since his results often differ essentially from mine, even as to sign, and since this is a master of considerable importance for theoretical considerations, some critical survey of his results seems called for. In general, Lussana finds that the thermal conductivity of all metals increases under pressure, and this increase is nearly the same as that of the electrical conductivity, so that the Wiedemann-Franz ratio remains nearly constant under changes of pressure. Lussana's method was an adaptation to high pressures of one origi- nally due to Depretz and Biot. A long bar of metal has a source of heat at one end and is immersed in a medium through which the heat may flow away laterally. The temperature of the bar, which is as- sumed constant across the section, is measured at three equi-distant points along it, and in terms of the two differences of temperature thus obtained, a relation can be found between a certain geometrical factor and the ratio between the thermal conductivity and the lateral con- ductivity into the surrounding medium. The essential difference between this method and mine is that in mine there is a source at one end of the bar and a sink at the other, so that nearly all the heat input EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 123 flows through the bar and out at the other end, and only a compara- tively small part is lost laterally to the surroundings; whereas with Lussana all the heat input flows out laterally. In Lussana's method the correction for the efl^ect of pressure on the transmitting medium affects directly the entire heat input, whereas in my method the pres- sure correction is to be applied only to that part of the heat input which escapes laterally. My most serious criticism of Lussana's method concerns this cor- rection for the transmitting medium. The magnitude of the correc- tion is about 30% per thousand kg., whereas the order of magnitude of the changes of thermal conductivity of the metals is at most only 3%, or one tenth of this. This demands that the effect of pressure on the transmitting medium be known ten times well as the final result for the metal. Nevertheless, Lussana determined the correction for the liquid to only one significant figure; as a matter of fact there is a mis- print in his paper, which made the correction appear to be at the rate of 300% for one thousand kg. I inquired about this in a letter to Lussana, and he told me that the decimal point had been displaced one figure, and that the correct result was 30%, agreeing with my own results as far as order of magnitude goes. Having determined the correction to one significant figure, not even noticing the departure of the effect from linearity with pressure, Lussana gives his coefficient for metals to three significant figures. Three significant figures for the metal would have demanded at least four significant figures in the correction. Lussana states that his results were computed from the observa- tions by the method of least squares ; he does not anywhere reproduce a single set of observations, nor does he state the probable error of his results, sui'ely a significant omission considering the method of computation. There is no clue in his paper to the accuracy to be attached to his results. There seems to be almost no correlation between Lussana's results and my own. In only one case, that of zinc, do we find the same sign for the change produced by pressure in the Wiedemann-Franz ratio. It seems to me that for the present we are justified in assuming that there are large errors in Lussana's results. Discussion. Probabl}' the most significant theoretical conclusions from the aljove data may be derived from the pressure coefficient of the Wiede- mann-Franz ratio. The classical electron theory would lead us to expect that the coefficient would be zero, since the ratio is the same for 124 BRIDGMAN. all metals at the same temperature, and the same metal under different pressures at the same temperature is merely a special case of two different metals. As a matter of fact the ratio is not constant, but may either increase or decrease with increasing pressure; in the ma- jority of cases it decreases. The average values of the coefficient between 0 and 12000 kg. are shown in Table III. TABLE III. Metal C'ocllicicril of W iedciiiaiin-Frunz Hatio Pb +O.O56 Sn +0.063 Cd -O.O5I7 Zn -0.062.1 Fe -O.O52O Pt -0.063.5 Ag -O.OsTO Cu -0.0693 Ni -o.oa3 Sb -O.O4IO Bi -O.OiK) My own theory of electrical conduction attempted to explain the | Wiedemann-Franz ratio,® and to do this, I imagined the .same sort of mechanism of conduction as the classical theory. I still can see no reason to suppose that the most important part of heat conduction is not as imagined by the classical theory; the success of the theory in accounting for the numerical value of the ratio, which is approximately constant for the different metals, (it may vary from 6.38 X 10^" for aluminum to 9.14 X 10'" for bismuth), is too striking to be put aside \ with no substitute. At the same time it is evident that the account given by the classical theory cannot be complete; no account has been taken of the conduction by the atoms, and the agreement of the theoretical with the experimental value is not as close as we must ' demand of a finished theory. It is natural to look to the still unexplained part of thermal con- ductivity to account for the departures from constancy of the ^Yiede- mann Franz ratio under pressure. The part of the conductivity EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 125 which is due to the electrons would be expected to have the same pressure coefficient as the electrical resistance (except for a possibil- ity to be mentioned later); the remaining part must be capable of either positive or negative variation under pressure, and must be of the right order of magnitude. In the first place, let us consider the possible magnitude of the non-electronic part of heat conduction. The first deduction of the theoretical value of the Wiedemann-Franz ratio, given by Drude, was an elementary one, in which certain simplifying assumptions were made, particularly that the velocities of all the electrons were the same. Later Lorentz gave a more exact discussion, taking account of the ^Maxwell distribution of velocities among the electrons, and ob- tained a value for the ratio only two thirds of that of Drude. The discussion of Lorentz has later been verified by Bohr and others. The elementary value for the ratio is much closer to the experimental values than the more rigorous one, but still lies somewhat low. The failure of the more exact value to agree more closely with the facts has been regarded by some as a blot on the classical theory, but by others is regarded rather as to the credit of the theory, because the Wiedemann-Franz ratio as calculated by the elementary theory was felt to be too close to the experimental value to sufficiently allow for the atomic part of the conduction. I shall take this latter point of view, and consider that the value for the Wiedemann-Franz ratio calculated by Lorentz represents the part due to electronic conduction, and that the dift'erence between this theoretical value and the actual value represents the part of the heat conduction that must be accounted for in other ways. This point of view at once imposes certain numerical limits on the changes under pressure that it should be possible to obtain experimentally. For the total change of thermal conductivity under any pressure must never be so great as to more than wipe out the part of the conductivity which was initially ascribed to the non-electronic part. This means that the total decrease of thermal conductivity, after allowing for a change equal to the change of electrical conductivity, must not be greater than the difference between the total initial thermal conductivity, and the part given by Lorentz's expression. In practise this imposes a restriction only when the thermal conductivity increases under pres- sure less rapidly than the electrical conductivity. An examination of the results obtained in this paper will show that this condition is met in all cases. The condition imposed is most restrictive in the case of nickel. Under 12000 kg. its thermal conductivity decreases by 14.5%, 126 BRIDGMAN. and the electrical conductivity increases by 1.8%. The sum of these, 16.3%, is an upper limit which the fractional part of the total con- ductivity due to the non-electronic part must not exceed. Now the theoretical value for the Wiedemann-Franz ratio is 4.3 X 10^'' (Lo- rentz's value), and the experimental value for nickel is 6.99 X lO^**. This allows the possibility of 39% of the thermal conductivity initially being of non-electronic origin, which is more than twice the extreme set experimentally. It would seem that under these conditions, when so comparatively large a part of the atomic conductivity has been wiped out by pressure, that the relation between conductivity and pressure must depart from linearity. The experimental accuracy was not great enough, however, to show such a departure. Further consideration of the theoretical significance of these results is reserved for a forthcoming paper in the Physical Review. Summary. Two methods are described for measuring the thermal conductivity of metals under pressure. The first of these is a radial flow method, which has many theoretical points of advantage, but is of limited applicability in practise because of the difficulty of getting metals in a condition of sufficient homogeneity. The second is a longitudinal flow method, the essential of which is the small size of the specimen. The irregularities of the individual readings by the second method are greater than by the first metliod, but the eft'ect of inhomogeneities is less and different specimens of the same metal will give the same result. Measurements of the effect of pressures to 12000 kg/cm^ on the thermal conductivity of 11 metals have been made by one or the other of these methods. The effect may be either positive or negative, and is more often negative than positive. In only two cases, lead and tin, does the Wiedemann-Franz ratio increase under pressure; for the other metals it decreases, and sometimes by large amounts. In addi- tion to the metals, the pressure coefficient of thermal conductivity of petroleum ether has been measured. The conductivity increases by a factor of about 2.2. The only previous measurements have been by Lussana, who ob- tained results entirely different from those found here. His method is criticised in some detail, chiefly on the basis of the uncertain correc- tion for the lateral loss of heat to the pressure transmitting medium. These results indicate that a fairly large part of thermal conduction EFFECT OF PRESSURE ON CONDUCTIVITY OF METALS. 127 in a metal is performed by the atoms. Theoretical reasons are given for estimating the atomic contribution to the thermal conductivity as 50 per cent of the electronic contribution. I am much indebted to the skill of my assistant Mr. J. C. Slater, who made a large number of the readings. It is also a pleasure to acknowledge my indebtedness to the Rumford Fund of the American Academy of Arts and Sciences for a grant with which a part of the expenses were defrayed. 1 P. W. Bridgman, Proc. Amer. Acad. 52, 571-646, 1917, and 56, 59-154, 1921. Phys. Rev. 9, 269-289, 1917, and 17, 161-194, 1921. 2 S. Lussana, Nuo. Cim. 15, 130-170, 1918. 3 C. Niven, Proc. Roy. Soc. 76, 34-48, 1905. 4 P. W. Bridgman, Proc. Amer. Acad. 53, 267-286, 1918. 5 G. Borelius und A. E. Lindh, Ann. d. Phys. 51, 607-620, 1916. 6 Reference 1, fourth paper. The Jefferson Physical Laboratory, Harvard University, Cambridge, Mass. VOLUME 56. 1. Kennelly, a. E., and Kurokawa, K. — Acoustic Impedance and its Measurement. pp. 1-42. February, 1921. $1.25. 2. Beix, Louis. — Ghosts and Oculars, pp. 43-58. February, 1921. $.85. 3. Bridc.man, p. W. — • Electrical Resistance under Pressure, including certain liquid Metals, pp. 59-154. February, 1921. $1.25. 4. 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Introduction 1^1 Historical Survey ]^^ Outline of Method 1^5 Experimental Details |42 Dimensional Discussion of the Cooling Process 149 Various Experimental Checks and Precautions 153 Experimental Procedure and Data 159 Other Possible Explanations of the Effect 164 Computation of the Departure from Ohm's Law 166 Theoretical Discussion 168 Summary ^'^ Introduction. It is to be expected that at high current densities the usual Hnear relation between current and E.M.F. in a metal will break down, that is, that Ohm's law will fail. J. J. Thomson,^ for instance, has shown that on the basis of the classical free electron theory of metallic conduction the current will eventually increase only as the square root of the E.M.F. at very high values, which means that the resistance will increase at high current densities. On the usual assumptions the current densities at which this effect will become important are of the order of 10'^ amp/cm^. Many attempts have been made to detect the existence of this effect experimentally, but hitherto without suc- cess. The chief obstacle to success is that the changes of resistance due to heating by the heavy current are sufficient to mask the changes of resistance due to a possible departure from Ohm's law. By the employment of a new method of attack, which avoids errors due to temperature rise, I have, I believe, not only succeeded in establishing the existence of the effect, but in measuring the departures from Ohm's law with some exactness in gold and silver. These results are described in the following paper. I find a departure from Ohm's law in the direction of an increase of resistance at high current densities, the maximum effect being of the order of 1% at a current density of 5 X 10*' amp/cm-. 132 BRIDGMAN, Historical Survey. A few attempts were made to detect the existence of the effect before Maxwell, but we may pass these over as not approaching in sensitiveness the method of ^laxwell.^ ^Maxwell was the chairman of a committee, the other members of which were J. D. Everett and A. Schuster, appointed by the British Association in 1876, to investi- gate the accuracy of Ohm's law. Two experimental methods were used, both apparently proposed by Maxwell; the experiments were performed by Chrystal. The first was a substitution method, by which the resistances of five similar coils were compared in various combinations of two in parallel and two in series against a single one. The current density in the single coil was thus double that in the multiple arrangement, and if there is an effect of the kind sought, the resistances should be different. A small positive result was found, but was ascribed to errors in view of the fact that the second method, which was much more delicate, gave negative results. The current densities employed in the first method were so low as not to cause apprecial)le heating of the wires. The second method was such that currents large enough to heat the wires to incandescence could be used. A heavy and weaker current were passed alternately in rapid succession through a fine wire, which was made one of the arms of an ordinary bridge. The period of alternation was so high that there were no appreciable cooling effects. Observations were made with a galvanometer of period long compared with that of the alternations. The apparent resistance was read first with the large and the small currents flowing in the same direction, and then with the small current reversed. Let us suppose that there is an effect of the kind sought, which means that the resistance for the large and the small current is not the same. If the galvanometer indicates balance, it must be because it is really off balance for both currents, to the one side for the large current, and to the other side for the small current. If now the small current is reversed the galvanometer will be off balance to the same side for both currents, and there will be a steady deflection. Hence if there is an effect, the balance will be altered by changing the direction of the small current. There were difficulties in the method. The period of alternation had to be chosen as high as 60 per second in order to avoid appreciable cooling effects in the short intervals of time when the small current replaces the larger one. There were consid- erable mechanical difficulties in designing an alternator of the requisite I FAILURE OF OHM's LAW AT HIGH CURRENT DENSITIES. 133 constancy, for it is obvious that the durations of the large and the small currents must be absolutely constant. A platinum contact dipping into a mercury cup and driven by a tuning fork was used, but always gave trouble, and the limits of accuracy were set by this part of the apparatus. The conclusion as usually quoted which was drawn from these experiments was that any deviation from Ohm's law must be less than one part in 10^^. This statement needs some expansion; it is obvious that no measurements can be made directly to this degree of accuracy. For one thing, changes of temperature of the surroundings absolutely preclude the direct attainment of any such accuracy as this. The meaning of the statement is as follows. Maxwell remarked that any departure from Ohm's law must involve only even powers of the current ; it is obvious that the first power cannot enter, for if so there will be a dependence of resistance on the direction of the current, which cannot be the case in an isotropic material. The initial departures from the law may be supposed to be proportional to the square of the current density. The maximum current density employed by Max- well was 5.6 X lO"* amp/cm^. At this density the resistance was found to have changed by not as much as 0.3%. Assuming the square law, this means that at a current density of 1 amp/cm- the departure from Ohm's law cannot be more than 1 part in 10^^. The original paper contains the careful statement of the conclusion in this form. The metals used by Maxwell were cylindrical wires of platinum (0.042 mm. diameter), German silver (0.051 mm.) and iron (0.14 mm.). The maximum current densities were respectively 3.4, 1.2, and 5.6 X 10* amp/cm2. Recently Wenner ^ of the Bureau of Standards has objected to the second form of Maxwell's experiment. He has repeated a modifica- tion of the first method with very much higher accuracy, and finds no deviation of as much as 3 parts in 10^. His objection to the second experiment is that negative results might be obtained even if there is a departure from Ohm's law. Thus if the potential difference across each arm of the bridge is proportional to the square of the current flowing through it, negative results will be obtained, because the bridge will stay in balance for any current, large or small. More generally, negative results will be found if the potential difference across each arm is the same function of the current for each arm. It is to be noticed, however, that this is not the manner of departure from Ohm's law which is to be expected. The departure sought for is not a function of the total current flowing through the resistance, but 134 BRIDGMAX. is a function of the current density. Furthermore, this function is known to be nearly independent of current density for low values. Doubling the total current in a conductor of small section will produce a much greater departure from Ohm's law than doubling the same current in a conductor of larger section. This is the only sort of departure from Ohm's law which we are looking for, or indeed which seems at all likely, and in my opinion Maxwell's experiment is entirely competent to answer this question up to its limits of error. As regards the other sorts of departures from Ohm's law, I believe that Wenner's position is sound. Since ^Maxwell, very few attempts have been made to detect the effect, probably because of the appalling sensitiveness of ^Maxsvell's experiment as usually quoted. Lecher * in 1906 made measurements on platinum and silver wires. He attempted to correct for the tem- perature effect in the platinum by observing the thermal expansion of the wire when carrying a very heavy current, and comparing the resistance to this heav\' current with the resistance to a feeble current passing through the wire when heated artificially to have the same thermal expansion as when carrying the heavy current. The diameter of the wire was 1 mm., and the current density was 3.8 X 10^ amp/cm-; there was no effect greater than 0.1%. The silver wire was 0.03 mm. in diameter. It was placed in a rapid stream of water, and the resistance measured under a current increasing until it fused. No temperature correction was applied. The apparent temperature when the wire burned out, using the ordinary temperature coefficient of resistance, and assuming that Ohm's law is true, was 130°. It is to be expected that this temperature would be somewhere between 100° and the melting point of silver. Hence within limits of error which may be several hundred percent, the experiment is consistent with Ohm's law. The accuracy is very much less than that of Maxwell; but on the other hand, the current density is ver\- much higher, reaching a maximum of 1.4 X 10^ amp/ cm'-, 25 fold greater. H. Rausch von Traubenberg ^ has attempted to avoid the tempera- ture difficulty by employing a condenser discharge of very short duration. The current densities that may be reached are higher than previously realized, attaining in one case a maximum of 10^ amp/cm"-. But the measurements of potential are inaccurate, being estimated by the break-down of a spark gap in air, and there are other sources of error arising from distributed capacities and inductances and the necessity of a long range extrapolation. I estimate that the error may certainly be as high as 10 or 15%. Within these limits, no effect was FAILURE OF OHM S LAW AT HIGH CURRENT DENSITIES. 13; found. The material used was constantan, of a diameter of 0.1 mm. The maximum current density reached produced a potential gradient in the wire of 400 volts per cm. With "Kruppin" wire, a gradient of 1500 volts/cm. was reached, although with not so high a current density. In spite of the very considerable error, these experiments have pushed the boundary of the validity of Ohm's law considerably farther back than the previous limit. "With regard to the materials used, it should be said that the maximum departure from the law is to be expected in those materials which, other things being equal, have a long free electronic path, and which are presumably the best con- ductors. From this point of view the most promising place to look for the effect is in silver, copper, and gold. Of course, on the other hand, it is to be said that in poor conductors it is possible to reach much higher potential gradients for the same current density, so that this advantage may outweigh the disadvantage. Outline of Method. In the method which I have used, the specimen exposed to the high current density is made one arm of a bridge, and its resistance is meas- FiGURE 1. Skeleton of the bridge connections. ured simultaneously for a heavy direct current and a small superposed sinusoidal current of acoustical fre- quency. If there is a deviation from Ohm's law the resistances to the two currents will not be the same. The following considerations show why this is. It must in the first place be remembered that a bridge is an instrument for testing the equality of potential of two points in a net work. In Figure 1, x denotes the arm of the bridge which contains the fine wire in which the Figure 2. Hypothetical re- lation between current (I) and e.m.f. (E) not satisfying Ohm's law. 136 BKIDGMAN. current density is to be high. In this branch we suppose that Ohm's law does not hold, but the relation between current and E.M.F. is given by a curve of the form shown in Figure 2. The other arms of the bridge, Ro, R3, and Ri are made of larger wire, in which the current density is always small, and hence their resistance is ohmic. Assume for the moment that it is possible to balance the bridge simultaneously for D.C. and A.C. Now let a heavy direct current h flow through z and Ro and a direct current I3 through R3 and Ri. Also let a small alternating current h sin w/ flow through x and R2 and is sin w< flow through R3 and Ri. The difference of potential between the ends of X is Ii tan 6 + ?'i tan 6' sin wt The potential difference across the galvanometer is (/i tan 6 + i\ tan 6' sin ut) — {I3 + is sin oi)t) R3. Since the extremities of Ro and Ri join, their potentials are equal, or /i (tan 0 + i?2) + h (tan 6' + R.2) sin o^t = {1 3 + (3 sin w^ {R3 + ^4). This splits into two equations 7i tan 6+ hR2= I3 (i?3 + Ri) (1) ii tan d' + h /?2 = is (Rs + Ri) (2) Now if the galvanometer is balanced for D.C. the constant part of the potential difference across its terminals must vanish, or 7i tan 0 - /a i?3 = 0. Combined with (1) above we get h R2 - hRi= 0, or, dividing tan d R3 Ro Ri (3) In the same way, if the bridge is balanced for A.C. the alternating part of the potential difference across the galvanometer vanishes, or ii tan 6' = is R3 Combined with (2) ll XL2 ^ I3 Ri and dividing tan e' R3 XV2 Ri (4) FAILURE OF OHM's LAW AT HIGH CURRENT DENSITIES. 137 ' Now conditions (3) and (4) are incompatible unless tan 6' = tan 6. Except for singular points, this means that the relation between current and E.M.F. in the arm x must be linear, or Ohm's law is satis- fied. Conversely if Ohm's law is not satisfied, the setting for balance will not be the same for D.C. and small A.C., tan 6 may be called the direct current resistance, and tan 6' the alternating resistance. They may both be determined by the ordinary bridge formulas by first adjusting i?3 and Ri for D.C. balance, and then readjusting them for A.C. balance. The departure from Ohm's law at a given current density, which I denote by D, is the fractional dift'erence between tan 6 and tan ^o, the tangent to the curve at the origin, that is, the re- sistance under small currents. This definition gives the equation for D : tan 6 — tan ^o tan ^0 It is now obvious that if we measure 6 and 6' at all points of the curve we can find the curve itself by an integration, hence the tangent at the origin, and so the departure from Ohm's law at any given current density. The mathematical details of this deduction will be given later. It is evident that the method in simple outline, as given above, avoids the difficulty of the unknown temperature correction because both currents are fiowing simultaneously, and hence the temperature of the wire is the same to both. There is, however, a temperature effect of a different kind from that usually met in this sort of experiment which arises as follows. The total rate of heat input under the cur- rent is proportional to the square of the total current, that is to (/i + ii sin oj/)-. The 27i ii sin cot term in this expression denotes an alternate heating and cooling, so that superposed on the large steady temperature increase there is a small sinusoidal fluctuation of temperature whose average is zero. But this small fluctuation of temperature produces a small fluctuation of resistance, and a heavy current flowing through a fluctuating resistance gives rise to a fluctu- ating difference of potential at the terminals of the resistance. There is, therefore, effectively introduced into the .r arm of the bridge a spurious additional sinusoidal E.M.F. which changes the A.C. balance. The action is similar to that of a microphone. We now discuss mathematically this spurious E.M.F. and the experi- mental means taken to eliminate its effects. We are for the present concerned solely with this effect, and in the following treat the resist- ance as ohmic. Any residual effect left after the elimination of this 138 BRIDGMAN. "microphone action" constitutes the departure from Ohm's law for which we are searching. Various tacit assumptions will be made in the course of this discussion which will be justified later. Return to Figure 1 for the bridge, and consider the heating effect in the arm x, treating its resistance as ohmic. The heat input is pro- portional to (/i + ^1 cos caty, where ii is small compared with 7i. Expanding this, neglecting the term in ij^ the rate of heat input is proportional to /i^ + 2/i ii cos cot, that is, there is a constant input proportional to 7i^, independent of the presence of the A.C., and there is a sinusoidal heating and cooling of the same period as the A.C. which is proportional in intensity to both the D.C. and the A.C. Under this heat input the conductor experiences a change of tem- perature, which may be analyzed into a constant change dependent only on the D.C, and a small alternating rise and fall, of the same period as the A.C, but not necessarily in phase wath it. The factor of proportionality which determines the amplitude of the alternating part is not the same as that which determines the amplitude of the steady part, but is a function of the period, becoming less for higher fre- quencies. Let us call the stead^^ change of temperature to, the ampli- tude of the in-phase part of the alternating part ri, and that of the out-of-phase part to. If the heat input is removed rapidly, t^ will be small compared with ri. The increase of temperature above that of the surroundings is therefore to + n cos cot -\- t2 sin cot. Now if Rq is the initial resistance at the temperature of the surroundings, a the temperature coefficient of resistance, and R the actual resistance when the current is passing, we have R = i?o [l + a (to + Ti cos cot + T2 sin wt)]. The potential difference across the terminals of x is R{li + ii cosoj^). Expanding this by substituting the value of R above, and using the relations cos^ 0 = i (1 + cos 26), 2 sin 6 cos 6= sin 2d, we get : Potential difference = i?o( A(l + arc) + I hcxTi} -f RolhaTi + fi(l + q:to)} COS wt + Ro{liaT2} sin cot + i?ol| ^i«Ti| cos 2cof + Ro\^ iiocTo} sin 2cot. FAILURE OF OHM's LAW AT HIGH CURRENT DENSITIES. 139 Using the same notation as before for the current in the other arms of the bridge, we have at D.C. balance and Roh il-\-aTo-\-^aTij hRo = hRi. = ^3 ^3, Dividing to ehminate the currents, we obtain i?fl hi 1 + ocTo + lari - r = jRo Rs This shows that in general, even neglecting the cos^ term in the heat input as we have above, the D.C. balance will depend on the A.C. But this effect is doubly small, since ri is small compared with to and ii small compared with /i, and hence the effect may be neglected. The correctness of this assumption was checked experimentally. \Yith regard to the A.C, the expression above shows that there cannot be complete balance. There will always be higher harmonics, and there will be an out-of-phase component (in sin co<). These terms are small, as examination of the coefficients shows, but may neverthe- less be perceptible. The ear can set on the fundamental alone, and so eliminate the higher harmonics. The out-of-phase component gives rise to a smearing out of the sharpness of the minimum. This can be corrected by introducing another out-of-phase component to neutral- ize it by a variable mutual inductance between input and detecting circuits. The equilibrium conditions for the in-phase component are Roil and Eliminating the current, R, = is Rs i\Ri — iz Ri- h' 1 + a I To + n T V '^1 i?2 Rz Rd The condition for A.C. balance is therefore different from that for D.C. balance, the large term Ii/ii occurring in the expression for A.C. balance against the small term ii/Ii in the expression for D.C. balance. As the experiment was actually performed, Ro was kept constant, and Rs and Rt were varied. R3 and Ri consisted of extension coils 140 BRIDGMAN, connected to a bridfje wire, wliich was tapped by a moving slider. Hence adjustment was made by adding to R3 an appropriate resistance, and at the same time subtracting the same resistance from Ri. Let us call the initial resistance for balance with no heating effect (small D.C. or A.C. only) R^ and Ri. To maintain balance under the heavy D.C. with no A.C, R3 must be increased by AR and Ri dimin- ished by AR. A.C. balance with the heavy D.C. flowing is now main- tained by an additional increase of AR' to R3 and decrease of Ri by AR'. The conditions for balance under these three states of current flow are: Ro = R2 ^ (5) lU U. R. I + a = R-2 Rs + AR Ri-AR Ro\l+a( To + n '1/ R-2 R3 + AR + AR' Ri - AR - AR' (6) (7) Subtracting (6) from (7) and discarding squares and products of AR and AR' gives RoCtTi 1} _ 1 II Ri AR'iRs + Ri) /?4- - Ri {^AR + AR') Also neglecting hii/h compared with Ij/ii, and substituting for Ro its value from (5) gives Ri^ "^' /i Ri- (AR' -\-2ARy which gives again approximately From (6), for the D.C. setting, we get approximately Hence finallv we have a- '0 — AR ik 1 ^ R. r\ ii AR' To — h AR' (8) (9) (10) FAILURE OF OHM'.S LAW AT HIGH CURRENT DENSITIES. 141 If now the alternations are slow, so that at every moment the wire is approximately in thermal equilibrium, the factor of proportionality connecting ri with the alternating heat input is the same as that con- necting To with the steady heat input, so that we would have To = const Ii~ R } Ti = const 2/iti R ) which gives, substituting above in (10), AR' AR = 2. That is, for slow alternations, the difference between A.C. and D.C. settings due to the microphone action alone is twice the D.C. shift due to temperature rise under the steady current, and this relation holds no matter how feeble the alternating current. Since the steady rise of temperature is high, because the current density has to be pushed to the limit that the conductor will carry without burning out, it is obvious that at slow alternations the microphone action will entirely mask any sought for deviation from Ohm's law. The acoustical frequencies used in these experiments were not low enough to reach the extreme value 2 for the ratio AR' /AR. At the lowest frequency, 320 cycles, the ratio had reached about 1.2. At rapid rates of alternation, however, the conditions of heat transfer change. At low frequencies the thermal conductivity of the surroundings alone determines the equililjrium; at higher frequencies part of the heat input is used in raising the temperature of the sur- roundings and a term enters proportional to the .specific heat, and at still higher frequencies this term preponderates, and the factor of proportionality between amplitude of rate of heat input and amplitude of temperature alternation becomes proportional to the specific heat and inversely proportional to the frequency. We shall later apply a dimensional analysis to obtain more information about to and n, but for the present we may write, for any frequency and as before This now gives n = const /(oj) 21 ill R, To = const I^ R. AR' AR 2/(a,). 142 BRIDGMAN. At high frequencies /(w) o^ — , so that at high frequencies the micro- co phone effect is proportional to the reciprocal of the frequency, and it may be eHminated by proceeding to infinite frequencies (or zero reciprocal frequency). The procedure suggested by this analysis was that followed in the experiment. For a fixed D.C. the difference between the settings at D.C. and A.C. balance (that is AR') was observed over a range of frequencies, AR' was plotted against the reciprocal of frequency, and extrapolated to zero. The residual, if there is one, is the effect due to deviation from Ohm's law at the particular D.C. density in question. This procedure was repeated for a number of currents, and so the departure from Ohm's law obtained as a function of current density. Before proceeding further with the theoretical discussion it will pay now to describe the experimental details, in order that we may have an idea of the order of magnitude of the quantities involved. Experimental Details. The bridge was an ordinary four gap alternating current bridge, so constructed that inductive and capacity effects in the bridge were negligible. The resistance Ro, which was kept constant during the measurements on any single specimen, was a coil of heavy manganin wire immersed in an oil bath to carry away the Joulean heat. This resistance was approximately ec^ual in magnitude to the resistance x which carried the high current density. The resistances R3 and Ri consisted of heavy manganin coils connected by a slide wire, which was tapped by a slider. The wire was about 1 meter long, with a total resistance of about 3 ohms. The resistance of the extension coils was five or ten times greater than that of the specimen x, and the genera- tion of heat in them was so small that it w as not necessary to immerse them in an oil bath. The method of connecting the D.C. and the A.C. sources and of tapping across with the detectors for D.C. and A.C. balance is shown in Figure 3. The direct and alternating current sources are connected to the same terminals of the bridge, with a large inductance L in the D.C. line to prevent the A.C. backing into the battery, and a large condenser C in the A.C. line to prevent the D.C. backing into the A.C. source. D.C. balance was shown by a Leeds and Northrup high sensitivity galvanometer of about 8 ohms internal resistance connected FAILURE OF OHM S LAW AT HIGH CURRENT DENSITIES. 143 as shown with a high resistance i?7 in series and another small resist- ance in shunt to cut down the sensitivity. In the latter part of this work this galvanometer was replaced by another of less sensitiveness. The A.C. detector was a telephone tapped between the same points as the D.C. detector, but with a large condenser in series to prevent D.C. getting into the telephone circuit. The telephone was tapped across a transformer placed in this circuit. In this circuit is also one of the coils of a mutual inductance, M, the other coil of which is in series with the A.C. source, and is not shown. This makes possible FiGUEE 3. Details of the bridge connections. the elimination of the out-of-phase component by suitable adjustment. The A.C. was prevented from entering the galvanometer circuit by the high resistance in series with it, and by an open key when the galvanometer was not in use. The condenser in the telephone circuit proved an unnecessary precaution, the resistance of the transformer and mutual inductance being sufficient to prevent enough diversion of the D.C. into the telephone line to introduce appreciable error. The condenser was used in most of the work, but in some of the later read- ings it was omitted. The telephone was one of 1100 ohms resistance, made by the Western Electric Co., type 509 W. The transformer was one of the small ones of the General Radio Co. made for this purpose, type 166. In Figure 3, the resistances R5 and Re which are connected to the same points as the sources of the current constitute an auxiliary bridge. 144 BRIDGMAX. The intermediate point was put to ground. This is the regular method of avoiding capacity effects in the telephone. i?5 and Re were so large that there was no serious diversion of current from the bridge. The D.C. source was a storage battery connected in series with a General Electric Co. ballast lamp (iron filament in hydrogen) of 1.4 amp. capacity. In series with the lamp was a commercial ammeter with which the constancy of the input current was checked. Any desired fraction of the output of the battery could be diverted from the bridge by a variable shunt between the lamp and the bridge. The actual current into the bridge or through the sample was not measured directly, but was computed from the ammeter reading, and the resist- ances, which were measured with the requisite accuracy. If a heavier current than 1.4 amp. were needed, two ballast lamps could be used in parallel. The dimensions of the sample were such that in almost all cases the maximum current that it could carry without burning out was not over 1 amp. The source of A.C. was a vacuum tube oscillator. The methods of connecting this were the canonical methods, and need not be gone into here. I am much indebted to Professor L. E. Chaffee and Mr. S. Ballantine for assistance and advice in setting up this circuit. For the first readings a Western Electric Co., hot lime transmitting tube Type VT2 was used, but this soon was burned out, and for most of the work a G. E. transmitting tube, type T Pliotron, was used. Not only does the present differ from preceding attempts in the method of measurement, but also in the form and dimensions given to the metallic resistance carrying the high current density. In all preceding work the metal has been in the form of a fine wire, of diameter of the order of 0.001 inch or more. An elementary discus- sion will show that a wire of these dimensions will carry only a limited current of the order of lO'' amp/cm- in the most favorable case. The limit is reached at a rate of heat input so high that the interior of the wire is at the melting point while the outer surface is at 0°, the thermal conductivity of the metal just sufficing to carry off the heat input under the temperature gradient so produced. It is possible to gain somewhat by rolling the wire flat as a galvanometer suspension, but not a great deal. An elementary dimensional discussion will show that the only way to gain on the upper limit of current density is by decreasing the thickness of the specimen, so that a given difference of temperature between interior and exterior will give a larger tempera- ture gradient, and therefore a greater heat dissipation. The thinnest metal that can be obtained is in the form of beaten leaf, and it was FAILURE OF OHM's LAW AT HIGH CURRENT DENSITIES. 145 with gold and silver leaf that I made the measurements. I was afraid to try films deposited by cathode spattering because it did not seem to me that the condition of the metal was sufficiently like that of ordinary metals, whereas the leaf may be supposed to be more like the massive aggregates of metal of ordinary dimensions. However, a few experiments at the end with spattered films of gold gave the same results as beaten leaf of the same thickness, and my fears are probably ill founded. The next work that suggests itself in this connection is an extension of the results found here for silver and gold to other metals, using spattered films. The thickness of the gold and silver leaf was determined by weighing a known area, assuming in the calculation that the density is the same as that of ordinary metal. The thickness of the silver leaf was 2.0 X 10~^ cm. Three thicknesses of gold were used, 8 X 10~®, 1.67 X 10"^ and 5 X 10-^. It was with some difficultv that I obtained the intermediate thick- ness of gold. Gold is beaten out in comparatively large quantities at a time in books of gold beaters skin, a great many thicknesses together. The last stage of the beating reduces the thickness by a factor of 6, from 5 X 10~^ to 8 X 10~^, and only these thicknesses can be obtained commercially. I am indebted to ISIr. Drew of Province Court, Boston, for his kindness in interrupting the last stage of the beating, and at some trouble removing a few of the partially beaten leaves from a large book. The sheets so obtained from the partially completed process were not nearly as perfect as those from the nor- mally completed beating. The state of the metal in a thin film differs in some unknown respects from that in larger masses. It has long been known that the specific resistance of spattered films is several times higher than that of the massive metal. ^ The specific resistance of gold films has been shown to be very high for very small thicknesses, to decrease rapidly as the thickness increases up to a certain point where the resistance is about five times normal, from here on to remain nearly constant over a range of thickness of about 20 fold, and beyond this point to decrease to the normal value. The temperature coefficient of resist- ance of spattered films has been frequently observed to be negative. The films of leaf metal used in this work did not show such great abnormalities as the usual spattered films, but nevertheless the resistance was very different from that of the massive metal. The temperature coefficient of my gold leaf was about 0.0015 between 0° and 30°, and was the same for the two thicknesses with which most of 146 BRIDGMAN. the measurements were made, namely 8 X 10"® and 1.67 X 10~^ cm. The temperature coefficient of the silver was much more nearly normal, and was 0.0032, its thickness being 2.0 X 10"^. The specific resistance of these metals was much higher than normal. That of the thinnest gold varied from 8.4 to 19.7 X 10"^ ohms per cm. cube, average 11.6. The spattered films of the same thickness varied from 15.4 to 2.3.8 X 10~^ average 19.2. The normal value for massive gold is 2.42 X 10~®. The thicker gold had a higher resistance than the thinner, varying from 9.75 to 18.5, average 13.3 X 10~^. The cause of the high specific resistance of the spattered gold is doubtless to be found partly in the lack of crystalline structure and perfect coherence, due to its manner of formation. The high resistance of the leaf metal, on the other hand, is doubtless in large part due to mechanical imperfections. Examina- tion of the thinnest leaf under a microscope shows a large number of folds and creases; it is practically impossible to spread the leaf on a surface so that it will lie smoothly in a single unwrinkled layer. The mechanical imperfections in the thicker gold were even greater than in the thinner, as already mentioned, doubtless partly due to the inter- ruption of the beating process at a disadvantageous stage. Two samples of gold 5 X 10"^ thick had resistances of 15.0 and 10.0 X 10~^ not essentially different from the other pieces. The specific resistance of the silver leaf varied from 3.5 to 5.1 X 10-^ average 4.1 X lO"*'. The normal value for silver is 1.63 X 10-^ It is seen that silver lies much closer to the normal than does gold. Under the microscope it too was full or minute imperfections, but of a different character from the gold. There were no folds, but a number of minute round perforations through the leaf. | It would doubtless have been most desirable if these experiments could have been performed on more massive samples with the normal electric constants, but the necessity of conducting away the heat seems absolutely to preclude such a possibility. The resistance has to be artificially cooled if current densities high enough to obtain an appreciable effect are to be reached, and the problem of mechanical support had to be solved. For this purpose the leaf metal was mounted on a piece of glass. The glass was covered with a very thin coat of insulating enamel by dipping it in a very dilute solution of the enamel in chloroform, the leaf metal was blown or otherwise spread over the surface, and was then baked at 210° until the enamel was hard. Gold or silver leaf so attached to the surface of glass is full of minute flaws, but by a search under the microscope parts can usually be found of sufficient homogeneity. The leaf was FAILURE OF OHm's LAW AT HIGH CURRENT DENSITIES. 147 cut SO as to leave a narrow isthmus of the shape shown in Figure 4. It is the isthmus that carries the high current density. Connection to the leaf on either side of the isthmus was by means of fine leads of copper caught to the leaf with a touch of solder. A special tool had to be made for cutting the isthmus. The point of a very fine needle Avas made to travel in any desired direction across the surface of the foil, scratching through to the glass, by an arrangement of two screws Figure 4. The isthmus form of the specimen. at right angles to each other. With this device, under the lens of a microscope, the isthmus could be cut to a high degree of precision. The dimensions of the isthmus varied somewhat from specimen to specimen, but the length was of the order of 1 mm. and the width of the order of 0.1 nun. The dimensions of each specimen were measured with a microscope. The specimen was cooled by a stream of water flowing across the isthmus at right angles to its length. This water was delivered from a small glass nozzle suitably held and directed. At first kerosene was used as a cooling liquid, in order to avoid danger of short circuit, but the cooling was not sufficiently rapid and the desired current densities could not be reached. I also tried currents of compressed air and hydrogen, with results very much inferior to those even for kerosene. In order to protect the specimen from the short circuiting action of the water, it was covered on the upper surface, except over the isthmus itself, with an additional coating of enamel. Any enamel on the isthmus itself is fatal. At first I used tap water, but this was too conducting. Ordinary distilled water, however, proved to be suffi- ciently insulating so that no short circuiting effects from it could be detected. After the distilled water hatl been used for some time slight irregularities began to appear due to increasing conductivity from miscellaneous impurities picked up from the air of the room; these irregularities could be made to disappear by replacing the water with fresh. It is necessary that the velocity of the cooling water be maintained constant. For small streams, a syphon arrangement was satisfactory, but for more rapid delivery the proper head was maintained by air 148 BRIDGMAN. pressure obtained from a large compressed air bottle, and was regulated to any desired value with a safety valve of special construction. In addition to the gold and silver leaf, I made attempts to detect the effect in manganin wire 0.001 inch thick rolled flat, and with Wollaston wire of platinum about 0.00006 inch thick. The attempt with manganin failed because the heating effects were too large, due to the dimension of the specimen. The attempt with platinum failed because of mechanical difficulties in mounting the wire and su)>jecting it to a stream of water. It is possible that with more pains it might be feasible to obtain results with platinum in this way. The thickness of the leaf metals used in this experiment was more than sufficiently small to ensure conduction of the Joulean heat developed by the heavy current without excessive rise of temperature. To illustrate the order of magnitudes involved let us consider an example. One specimen of silver that gave good results had the fol- lowing dimensions: Length 0.536 mm., width 0.072 mm., and thick- ness 2 X lO"'' cm. The maximum current before the specimen burned out was 0.745 amp., and the initial resistance was 1.30 ohms. The heat input into this specimen was therefore: rr . (.745)2X1.30 , Heat input = -tTo S^^ cal/sec. 4.18 = 0.173 gm cal/sec. This heat flows out through the area of one face, which is 0.0536 X 0.0072 = 3.87 X 10-* cm^. The heat outflow per unit area is therefore (0.173)7(3.87 X 10^) = 4.5 X 10- cal/sec cnr. Since the thermal conductivity of silver is approximately unity, the temperature gradi- ent required to drive this thermal stream is 4.5 X 10- degrees per cm. But the total thickness of the film is 2 X 10"^, so that the extreme temperatiu'e difference in the specimen between front and back face is 4.5 X 10-' X 2 X 10-» = 0.009°. It is of interest to compare the heat input with the heat capacity of the specimen. Its volume is 3.87 X IQ-^ X 2 X 10"^ = 7.8 X IQ-^cml Taking for the specific heat of silver 0.056, and the density as 10.5, we find the heat capacity to be 10.5 X 0.056 X 7.8 X 10-^ = 4.6 X 10"^. If there were no heat outflow the temperature would rise at the rate of (0.173)/(4.6 X 10-^) = 0.038 X 10^ = 38,000,000 degrees per second. The magnitude of the steady temperature rise actually observed in this specimen was about 50°, or 5000 times more than the mean rise of temperature required to procure conduction of the heat input out of the metal itself. It is ol)vious, therefore, that practically all the FAILURE OF OHM's LAW AT HIGH CURRENT DENSITIES. 149 r resistance to heat out flow is in the thin layer of cooHng water immedi- ately in contact with the surface of the metal. Our previous estimate of the maximum current density that a wire can carry must be cut down many fold. A 0.001 inch wire cannot carry 10'^ amp/cm^ under practical conditions. It still remains true under these new conditions, however, that the only change of dimensions of the specimen which will increase the maximum obtainable density is a decrease of diameter. Our numerical example shows that the body of the metal can be regarded as approximately at a single constant temperature, both for the steady rise of temperature and for the alternating fluctuations. In the mathematical discussion above it was assumed that the tem- perature of the metal could be specified by a single number; the numerical discussion just given constitutes the justification of this. Dimensional Discussion of the Cooling Process. In order to get further in our understanding of the phenomena we must now consider in some detail the steady and alternating changes of temperature ro and ri, remembering that practically all the resist- ance to heat outflow is in the cooling water. It is of course not possible to give an exact solution; the best that we can do is to give a dimen- sional discussion. Let us consider in the first place the ecjuations of heat transfer in a fluid that is in motion. The equations may be obtained by a slight generalization of the process by which the equa- tion of heat transfer is deduced for a medium at rest. Let us suppose that the medium is homogeneous except for temperature differences, that its specific heat per unit volume is c and its velocity of motion at any point v. Consider a small closed surface S at any point in the liquid. The rate of rise of temperature of the matter within this surface is the total heat input divided by the heat capacity. The heat input consists of two parts. The first is the ordinary conduction across the boundary, and is I I k — dS, where k is the thermal con- ductivity. This assumes that the velocity v is so small compared with the velocities of molecular motion within the liquid that the ordinary process of conduction takes place independent of the motion. The second part of the heat input is that which is convected, and is — f/cTPndS. From these two expressions we get the equation 150 BRIDGMAN. Applying Green's theorem to the surface S, transforming the surface to vokime integrals, using the condition that Div v = 0 because the liquid is to be considered as incompressible, and removing the integral sign, gives for the differential equation of heat transfer dr k ^ ^ , — - = - v-T — V • Grad r . dt c This equation applies at points inside the liquid. There will also be an equation to fix the boundary conditions. This equation is of the ordinary type, independent of the motion of the liquid, and is merely the statement that the heat input across the boundary is equal to the conductivity of the fluid multiplied by the normal temperature gradient. Apply these equations now to the present problem. If the motion of the liquid is not turbulent, and if the lines of flow are not altered l)y the heat input, then at the surface of the metal the liquid flows in planes parallel to the surface, the velocity increasing from the surface. The determination of the velocity distribution is a problem of hydro- dynamics, and involves the viscosity of the liquid and the variables which describe the mechanical roughness of the surface, but as far as we are interested in the problem the elements which enter our heat equations are determined if we can specify the velocity gradient at the surface. The other elements which enter the equation of heat trans- fer are the thermal conductivity of the liquid and its specific heat per unit volume. Subject now to the restrictions mentioned, we may make a dimen- sional analysis of the situation. Notice in the first place that since the flow of water is transverse to the specimen the rise of temperature etc. is independent of the length, provided only that the specimen is long enough for us to neglect end effects. We now enumerate the elements with which we are concerned and their dimensions. Name of Quantity Symbol Dimensional Formula Average rise of temperature Rate of heat input per unit length Velocity gradient in liquid Thermal conductivity of liquid Specific heat of liquid per unit volume Breadth of specimen Frequency' of impressed heat input T T Q // L-1 r-i 9 f-i k HL'^ r-i T-i c HL-' r-1 b L CO r-i FAILURE OF OHM's LAW AT HIGH CURRENT DENSITIES. 151 Consistently with our numerical discussion we have not tabulated the thermal properties nor the thickness of the specimen itself, since the effect of these is vanishingly small. We have now two cases to consider; first the steady temperature rise. The period of the impressed heat input does not enter, and we have to find all the dimensionless products of the first six quantities of the list above. Since there are four fundamental kinds of quantity (instead of unit quantity of heat H, we might have expressed heat in mechanical units, thus replacing // by M, with no change in the final result), and hence two dimensionless products. Inspection shows these products to be kr/Q and k/gch-. Hence the relation which we want may be expressed as urn where / is some unknown arbitrary function. This relation can be tested by experiment, and so some idea obtained of the correctness of the assumptions underlying the discussion. For instance, at constant rate of flow of cooling water, the above equation shows that the steady temperature rise should be proportional to the rate of heat input, or to /-. We can obtain an additional check for low rates of flow. For low rates, but not too low, it seems natural to assume that an impor- tant part of the rise of temperature is inversely proportional to the rate of flow, or inversely as g. This means that in / there is an important term which is the reciprocal of its argument, and we obtain as a partial expression • T = Const Q/gclr'. Some experimental information may be obtained here by varying the breadth of the sample at approximately equal rates of flow. That the average rise of temperature should be less for the greater breadth seems somewhat paradoxical, and affords a more drastic test than the proportionality of temperature rise to the rate of heat input. Now let us consider the alternating fluctuations of temperature. To distinguish from the steady case, and consistently with the previous notation, we denote the amplitude of the alternating heat input by Qi, and the amplitude of the alternating temperature change by n. We now have seven quantities, and hence three dimensionless products. The additional product, beside the two already obtained, is gr/co. The relation between the variables now takes the form 152 BRIDGMAN. where

Kl and silvor. Tho uiaximum ounvi\t donsitios woro about r> \ 10" «'up ^''^^'' '*''^^ *^^^' doviations fwm (.'•hni's law woiv of tho onlor of ono por ivi\t. If (l»o nuvhanism of c\Muluotion is a froo path mivhai\ism. thoso ri^sults prv^bably moan that tho froo patli is nmoh U^n^^T than snppvv*tHl on tho olassioal oUvti\>n thix»ry. It is a ploasnrt^ to aoki\owK\lct' nty indobtixlnoss to my assistant Mr. «1. C Slator for n\akinv: noarly all tho rt\»dinj::s. 1 J. .1. ri»oT\tsvM\. Tho iVrpusovUtjr Thtvrv vM Mattor. l^H^T. p. oo. a iMork MawvoU, .1, H, Kvon^tt. A. Sohvistor. Brit Asi:oo. Kop. 1S70. otH>3. S F. Wonuor. rUvj*. Kov. oJ^ lo. :>;ii >>;vj. UV.XV 4 K. l.n IVtuilvnlvrs:. rh\->!- Z8. IS. To-TS. H)17. e J. r.Httorv\>n. Phil. M,s»i. 4. tv\2-<>7S, UXVJ. : W. R Ot. 8\vann. ^hil.^l;^>i. JS. U>: 4VHv 1914. ^ r W". l^rivi^iitnan. rivv^ Kov, i>. Jl^iWJSi). 1917, and 17. UU-liM. 1921. H.srN~>;rvi ruivorsitv, 0;uubrids?\ Mas?, VOLL'ME 50. J. K p.wwr.fxy, A. K,, arid Kiiiumawa, K, - A«y»«wtic Inn/fAitiHA: hiiA it« Vf»!««M/'wn»!nt, f/(.. 1-42, VMiritarY, J'^21. »I,2/l, 2, V>f.i.t.,\ji>t>in.~('t\fmiMnwl()'-At\itr%. fr\>. VA-TA. VfAffuury.XWl. %JA. a, tU-ilf.viA-i, I', W . ~ yAfJ,Ui<:Hi U'9*iti.ittityi: tiu'ir.r Vtm^uiff., iwiuAUtK r/^Utiti I/qui/J M<-l:iU, j;j<. r/J J.'',4, F'-,»/fu.-.ry, i;»2), «).2./. 4. l.ifHA. Ji/HKi'H. - Sl'Aum ifit 'A Hiir1iii:K ti/r any t'*Mitiow>l Vif,\i\ '/ y*Tit'i'ln In \'Mi*tttiuiiniuxl! Kfi«r«y */ F'/f/H.-iti//n '/ Th.dl'/Mfi f'xli/J*;, w- H'? 2^'/. Aj/ril, r->2). 7 Ul-.llll'.l., W, A,— AfWiimafj/Wn I^^X^k 'n- i'.uiVw, \ Kf.<.-*r. C; M',. ini^., Wi\. $1.2.7. 9. Hrr'.n/xv;K, Fka-.k i- - 'rji* Ktm f. pp. a2'«^5J5J, Jufi*-, I'^l. $.75. 10, Ciw*«»«i, CHkhUVM H, — TJi*! Hiimfttrd Vitttf3nU f'<*f>»; MyUiVn* Hfni fttan/Jinjf V(/,) }. (5) Likewise, for h positive. Vl -jh= ± {f{h)-jg{h)}. (6) 176 PIERCE. 3. To Extract the Square Root of P -\- jU, where P and U are any Real Quantities. — Factoring out the square root of P, we have by (5) and (6) VP+jU = ± Vp Ifih) -hjg(h)}, if U/P is positive, = ± VP {fig) - j g{h)}, if U/P is negative, (7) where h = U/P. 4. To Find sinh-^ (P + jU) where P and U are Both Positive and Real. — Let smh-' {P +jU) = A+jB, (8) then P + jU = sinh A cos B -\- j cosh A sin B, whence P = sinh A cos P, (9) U = cosh yl sin B. (10) Regarding signs, in accordance with the caption, we see that, since cosh A is always positive, sin B is positive and sinh A and cos B are both positive or both negative. Taking the sura of the squares of (9) and (10), we have P2-I- u^= sinh^ A cos2 B + cosh^ A sin^ B = sinh2 A cos2 P + (1 + sinh^ A) (1 - cos- B) = 1 - cos^P + sinh2.4, whence by using (9), we have P2 OS' P + , „ , and COS" P (11) P2 ii-ii-t'' 1 (13) '"^^^^ ~ Sinh-^P- 1 - P2- f72 Letting V = V^^' (14) and solving (11) and (12) as quadratics, we obtain cos-p = r ± \/p2+ V% sinh^.l = - !'■ ± \/P-+ r^. TABLE FOR TRANSMISSION LINE PHENOMENA. 177 Using the proper signs before the radicals to make the expressions positive as demanded by their left hand equivalents, extracting the square roots and employing (1) and (2), we obtain the following results If V>0, sinh .4 = ± V2Vg{h), cos 5 = ± VWjQi), if r<0, sinh A = ± V- 2Vf{h), cos 5 = ± V- 2V g{h), where h = P/V. Having regard to the rule of signs enunciated under (10), we may WTite sinh-i (P + jU) = X + i (?/ + 2Trn) and - .r + i (tt - y + 27rw), (15) where if V >0, X = sinh-i { + V2F g{h) ],y= cos-i { +VWf{h) \ , (16) if F<0, X = sinh-i {+ V-2Vf{h)], y = cos-^+V- 2V g{h)],{ll) with h = P/V. (18) Equation {15) gives the value of sin-^ (P -\- jU), xohere P and U are real, positive quantities, in terms of x and y defined by {16) and {17). The value of V is given by {14). The signs in {15) are so chosen that the value of y in the first quadrant is to be employed. 5. To Find sinh-i (P - jU), where P and U are Positive, Real Quantities. — This differs from the preceding problem only in the fact that sin B is negative, whence s\nh-\P-jV) = x-j {y+2Trn) and -.r+j (i/+7r+27r70, (19) with y in the first quadrant and with x and y defined as in (16), (17) and (18). 6. To Find sinh-^ {-P-jU) and sinh-i (-P + yC/), where P and U are Positive, Real Quantities. — These results may be had directly from Sections 4 and 5 by use of the facts that and sinh-i (-P - jU) = - sinh-i (P + jU), (20) sinh-i (- P + jV) = - sinh-i (P - jU). (21) 178 PIERCE. 7. To Find cosh-i (P + jU), where P and U are Positive and Real. — Let cosh-HP -\-jU) = A -{-jB, (22) then P + jU = cosh A cos B -\- j sinh A sin jB, whence P = cosh /I cos B, (23) [7 = sinh A sin 5. (24) The sum of the squares of these two equations gives P2 -j- u^= cosh2 J cos2 B + sinh2 .4 sin- B = 1 + sinh- A — sin^ B, whence by substitution from (24) and by solution of the resulting quadratic equations we obtain sinhM = - r ±Vr2+ V\ and sin2 5= r±VL^2_|_pr2, These give, with choices of signs to make A and B real and satisfy (24) if V >0, sinh A = ^V2Vg(h), sin B = ^V2V f{h), h = U/V, if F<0, sinh .4 = ±V'-2r/(/0,sin5 = ^V~^2V g{h), h = U/V. In accordance with (23) and (24), in each line the sinh A and sin B have the same sign before their radicals, and the angle B must be so determined that cos B is positive. Whence cosh-i (P + jU) = ± {a + j {cp + 27rn), (25) where if F>0, a = sinh-M+V2r^(/0},

rg, a = Vlcc^^— rg g{h), /3 = V/cco- - rgfih), (37) rcco + gloi where h = - — - — ' — • Icco-— rg II. Ulcc^' re, ^.■=\'^^^l/W+i^(/OK (39) glw — row where h = ^^ , — - — ~. rg -\- lew IV. ligKrc ^i = \' '•'^' o'T \m - j g{h) } , (40) rcco — gl(j} where h = — ; — -. rg + lew 1 Obtained by introducing the /- and g- functions into familiar equations. Compare Kennelly: Applications of Hyperbolic Functions, pp. 70 and 125. University of London Press. TABLE FOR TRANSMISSION LINE PHENOMENA. 181 13. Constants of an Artificial Electric Line. — If 2i = complex impedance of the series elements of an artificial line, 22 = complex impedance of the shunt elements, M = mutual inductance (if any) between adjacent series elements, (p = retardation angle of current per section of line, a — real attenuation constant per section of line, and if w /e let and P+jU = V = 2l+ 2So 1 - r— r- (41) (42) where P and U are real quantities, then ^ I. IfV>0, _ _ a = sinh-V2F g{h),

0, a = sinh"^ V2f' dW^ ^ — sin h = U/V. -1 i V2vm} (6) with II. ifr0, ^=sin-i /^ /^/l+^ + l- (17) This equation is the equivalent of (142), p. 317, of Electrical Oscilla- tions and Electric Wares. \Ye now proceed to develop the subject further. Confining our attention to this case in which V is greater than zero (that is, A positive), and expanding the inner radical in (17), we obtain, after transposition and squaring, sinV= between Series Sec- tions of a Low Resistance Line. — Equation (7) is the general ex- pression for surge impedance of the line of Figure 1. When the series impedances are inductances and the shunt impedances are capacities, as in Figure 3, 2i and 22 take on the values given in (9) and (10). These substituted into (7) give 1 SvEt'- '" Equation {30Y is the general expression for surge impedance of a line of the type shown in Figure 3. In this equation L = L\-\- 2M, and Ro= VL/C'2. It is seen that, if the resistance of the line is low, the imaginary term in (30) is small, and the real term tends to approach independ- ence of CO as 4il/ approaches L. Making M = .lZi= L/12, as is required in order to make T less dependent on frequency, has the effect of cutting the real term containing or by about ^ and hence the introduction of M = .iLi reduces the dependence of surge impedance on frequency for low resistance lines. 1 The corresponding equation (142) p. 317 of EI. Osc. and Wares has in the first printing of the book an error, that has been corrected in the Second Im- pression of 1921. ARTIFICIAL ELECTRIC LINES WITH MUTUAL INDUCTANCE 209 III. An Artificial Line to Simulate an Actual Smooth Line. Improvement as to Attenuation and Surge Impedance brought ABOUT BY Proper Mutual Induction between Series Sections. 14. Constants for Actual Smooth Line. — In the case of an actual smooth line, let r, I, c = respectively the resistance, self-inductance, and capa- city per loop unit of length; a = real attenuation constant per unit of length; 13 = angle of lag of current per unit of length ; Zi= surge impedance of the line; then a = CO (3 = CO l\ 0 C /CO (31) (32) (33) These equations are the familiar expressions. See, for example El. Osc. and El. Waves, pp. 329 and 330. 15. To Design an Artificial Line with Lumped Sections that Will Simulate the Smooth Line as to Surge Impedance. — For this purpose we shall postulate a line of the type shown in Figure 3, and shall determine what value of M brings the surge impedance of this line, as is given in (30), most nearly into the form of (33) in so far as dependence of s,- on co is concerned. Since the real part of (30) is generally much larger than the imaginary part, it is seen by inspec- tion that a good approximation to this result is made by making M = L/4.; (34) which bv (13) means > M = Li/2. (35) 210 PIERCE. By making M have the vakie given in (34) equation (30), in view of the fact that ^0 = Vl/C., (36) becomes (37) Equation {37) gives the surge impedance of a lumpy artificial line with 31 = Li/3. Since the imaginary term in the surge impedance of a smooth-line {equation {33)) is generally small over practical ranges of frequency, it is seen that {37) is essentially of the form of {30) in so far as concerns dependence of surge impedance on frequency} 16. To Determine the Mutual Inductance between Series Section in a Lumpy Artificial Line to Bring it into Close Simi- larity with A Smooth Line as to Attenuation Constant.^ Postulating a line of the type of Figure 3, and substituting (9), (10), (13), (14) and (15) into the value of a given in (5) we obtain If F>0, \~i i I r (38) Equation {38) is the general equation for attenuation constant for a line of the type of Figure 3, tinder the condition T' > 0. Equation (38) expanded with neglect of higher powers of rf'/A^ gives smha=,^\j ^1-— +-- + •••• f (39) A corresponding expansion of (31) gives approximately for the smooth line « = fVfc.|l-f +■■•[, (40) 1 This fact was called to my attention by Mr. Phillip Machanik, who based his observation in an examination of the equations in Electric Oscillations and Electric Waves. \ ARTIFICIAL ELECTRIC LINES WITH MUTUAL INDUCTANCE. 211 where 770 = r/Zio. (41) Since the second terms in (40) and (39) are usually small, and since a is also sufficiently small to make sinh a essentially equal to a, the two equations reduce to nearly the same form in respect to co if Q/A = iCaco^. (42) Replacing A in (42) by its value from (14) with neglect of 77- in (15) w^e obtain ^ " 1 + i>cwy4' which compared with (14) shows that (42) is approximately satisfied when M = L/4: = ii/2. (42) A substitution of this value of M into the exact equation (38) gives in a careful approximation sinh a = I Via, CO I 1 - f + ^ (iCoV + I U~C,w) I . (43) If now w^e expand the hyperbolic sine into sinh a = a -\- a^/6, and replace the a in a^ by the first term of (43), we obtain a = IVLC, CO I 1 - 3 + ^ (lC^co^ + I UC.}u:) \ ■ (44) The approximation of equation (44) shows that even when rj. is as large as . 5 and ivith LCoui^ as large as 1 the introduction of a mutual inductance of the value given in {4'2) makes the attenuation constant of the artificial lumpy line of the same form as the attenuation constant a of the smooth line, and that the two attenuation constants can be thus made to agree over a wide range of frequencies. 212 PIERCE. IV. Conclusions. The following results, believed to be novel, for artificial line con- struction have been here derived. 1. To obtain a minimum dependence of time lag per section on frequency of an electric artificial line of low resistance, the line should be constructed with mutual inductance between neighboring series inductive elements equal to one-tenth of the self inductance of each series inductive element. 2. To most closely simulate a real smooth electric line as to attenu- ation constant and surge impedance by the use of an artificial electric line with lumpy sections, there should be in the artificial line a mutual inductance between adjacent loops equal to one-half of the series self inductance per loop. 3. Details for calculating the performance of lines constructed according to 1 . and 2 . are given. Cruft Laboratory, Harvard University, Cambridge, Mass. VOLUME 56. 1. Kennelly, a. E., and Kurokawa, K. — Acoustic Impedance and its Measurement. pp. 1-42. February, 1921. $1.2.5. 2. Bell, Louis. — Ghosts and Oculars, pp. 43-58. February, 1921. $.85. 3. BniDGMAN, P. 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THE PARASITIC W0R:MS OF THE ANIMALS OF BERMUDA. I. TREMATODES. By Franklin D. Barker. With Three Plates. CONTRIBUTIONS FROM THE BERMUDA BIOLOGICAL STATION FOR RESEARCH. NO. 135. THE PARASITIC \YORMS OF THE ANIMALS OF BERISIUDA. I. TREMATODES. Franklin D. Barker. Received Feb. 12, 1922. Presented by E. L. Mark, March 8, 1922. Introduction. Through the courtesy and generous assistance of Professor E. L. Mark, Director of the Bermuda Biological Station, and of the National Academy of Science, it has been my pri\dlege to spend two seasons at the Bermuda Station, collecting and studying the parasitic fauna. The following paper is the first of a series which will embody the results of these investigations. The two forms here described were found in the stomach of a Hawk's- bill Turtle, ChcJonia imbricata (Linn.), at the Bermuda Biological Station. Pachypsolus brachus, n. sp. (Pis. I and II, Figs. 1-8, 12). 1 . Morphology. General Appearance. — The description of the following species is based on the study of 27 preserved specimens, 1 1 of which were killed and fixed in 2% formol and 16 in vom Rath's osmio-sublimate mixture. Little difference can be seen as a result of the different killing reagents other than in color. Specimens fixed in formol are grayish-yellow, those fixed in vom Rath's fluid black. A detailed study has been made of specimens in toto, both unmounted and mounted, and of series of frontal and sagittal sections. The body is oval and plump (PI. II, Fig. 8), being one half as thick as wide. The length varies from 3jnm. to 3.7 mm., the mode being 3 mm., which is the length of 50 per cent of the individuals. The width varies from 1.5 to 1.9 mm., the mode being 1.7 mm., which is the width attained by 60 per cent. The ends are bluntly rounded, 216 BARKER. the anterior end slightly more tapering than the posterior. In the median line at the posterior end is a well defined terminal invagina- tion, which marks the position of the excretory pore. The dorsal surface of the body is strongly arched, the ventral surface slightly cupped. The sides are nearly parallel with the exception of a wide, shallow constriction midway between the ends at the level of the acetabulum. Spines or scales were not found anywhere on the body. In the preserved specimens, the anterior third of the body shows a marked and constant tendency to flex ventrad, which gives rise to a well defined and rather deep ventral cup between the oral and ventral suckers. This cup-like depression persists in compressed specimens and possibly functions as a secondary holdfast (PL II, Fig. 8). The oral sucker is comparatively large, well defined, nearly circular in outline and ventral in position, with its dorso-ventral axis at right angles to the chief axis of the body. In compressed specimens the oral sucker is 0.80 mm. to 0.82 mm. wide by 0.66 mm. to 0.82 mm. long. In frontal sections it measures 0.82 mm. wide by 0.82 mm. long. The ventral sucker, or acetabulum, lies in the median area at the posterior margin of the anterior half of the body and faces obliquely cephalad. It is of approximately the same size and shape as the oral sucker. It measures 0.60 mm. to 0.74 mm. in length by 0.70 mm. to 0.74 mm. in width in compressed specimens and 0.72 mm. by 0.80 mm. in frontal sections. The genital pore is not salient and lies in the median line, at the anterior margin of the acetabulum, or else slightly to the left of, and just anterior to the acetabulum. In the middle quarters of the body (PI. I, Fig. 2), along the sides, and extending well toward the median line on the dorsal surface, can be seen the characteristic dark colored, convoluted tubular and finely granular vitelline glands in moss-like patches. The uterus appears as a dark coiled mass nearly filling the \entral field of the posterior third of the body. Digestive System. — The transversely oval mouth leads into the triangular lumen of the oral sucker, which is 0.90 mm. deep in sagittal sections with thick muscular walls. A thick walled, large and power- ful cup-shape pharynx follows immediately (PL II, Fig. 12). The pharynx measures 0.58 mm. long by 0.52 mm. wide by 0.44 mm. deep in sagittal sections. Eight longitudinal muscular ridges or folds project from the inner' wall of the anterior two thirds of the pharynx into its lumen. Of the four larger or primary ridges, one is dorsal, one ventral, and two lateral; alternating with these are four smaller PAKASITIC WORMS BERMUDA. I. TREMATODES. 217 or secondary ridges. An oesophagus is not present, the common transverse caecum following immediately behind the pharynx. On each side of the pharynx a single well defined diverticulum extends cephalad from the transverse caecum lateral, or dorso-lateral, to the oral sucker to a height of one half the sucker's depth. These diver- ticula may or may not bifurcate at their terminations. The lateral caeca are broad and deep and they extend in an undulating course to the posterior end of the body, where they end blindly, giving off in their course numerous small lateral and deep ventral diverticula. The caeca are deeper than wide and lie in a plane mid-way between the dorsal and ventral surfaces in the lateral fields of the body (PI. II, Fig. 12). Male genitals. — The two testes (PI. I, Figs. 1, 5), of medium and nearly equal size, irregular in shape and with undulating to slightly lobed margins, are situated in the same transverse plane midway between the anterior and posterior ends of the body. The testes vary in size from O.oS mm. to 0.68 mm. by 0.34 mm. to 0.54 mm. The right one is slightly behind the posterior margin of the acetabulum and its left end projects beyond the median plane into the left half of the body. The bulk of the left testis is farther from the median plane than is the left margin of the acetalnilum, and its anterior end extends a little farther forward than the posterior margin of the acetabulum. From the antero-dorsal margin of each testis a small vas efferens passes obliquely forward and toward the median plane. These unite just cephalad and dorsad to the OA'ary to constitute the vas deferens (PI. I, Fig. 3); this continues in the median area to the base of the cirrus pouch, which it enters. The cirrus pouch, though compara- tively short, is an elongated pear shaped organ situated immediately anterior to the acetabulum, with its long axis nearly perpendicular to the frontal plane of the body. It is 0.96 mm. long and from 0.24 mm. to 0.2S mm. in diameter, I)eing a little longer than the acetabulum is deep, so that its base lies slightly dorsad and cephalad to the acetab- ulum. It is so short that it does not bend around the acetabulum. The vas deferens upon entering the base of the cirrus pouch vmites with the enlarged transversely coiled seminal vesicle, which fills the basal third of the pouch. The seminal vesicle connects, in turn, with a comparatively wide prostatic duct, which has an undulating course and tapers toward the distal end of the pouch, where it merges into the ejaculatory duct of the short cirrus. The cells forming the prostate gland (PI. I, Fig. 3) occupy the peripheral portion of the pouch and extend from the seminal vesicle 218 BARKER. nearly to the distal end of the pouch. Fine ducts leading from the prostate cells occupy the medullary portion of the pouch and enter the prostatic duct. The lumen of the cirrus is lined with cuticula, while the lumen of the prostatic duct is covered with high filamentous papillae. The Avail of the cirrus pouch possesses a heavy outer sheet of longi- tudinal muscle fibers and a thin inner one of circular fibers. The cirrus, which is approximately one-fifth the length of the cirrus pouch, has an outer and an inner muscular component. The outer compo- nent comprises an outer sheet of longitudinal muscle fibers and a heavier inner sheet of circular fibers. The inner muscular component immediately surrounds the lumen of the cirrus and the prostatic duct and is likewise composed of an inner sheet of circular muscle fibers and an outer sheet of longitudinal fibers. The cirrus pouch is anchored and possibly controlled by a pair of oblique muscles which are attached respectively to the cephalic and caudal faces of its base. The cirrus opens into a common genital atrium, which has its out- let in the genital pore lying in, or a little to the left of, the median line and slightly anterior to, or just under, the anterior margin of the acetabulum. Female genitals. — -The ovary (PI. I, Figs. 1, 5, 6, PI. II, Fig. 7) lies near the middle of the body, in the median area, dorsal to the posterior portion of the acetabulum and is from one-half to two-thirds the bulk of one of the testes, globular in general form with undulating or slightly lobed outline. In the specimen figured (Fig. 5) it measured 0.38 mm. by 0.38 mm. The oviduct leaves the ovary from the middle of its anterior margin and at once turns sharply mediad; after making several loops it passes caudad in descending transverse coils, lying a little to the right of the median plane, to the end of the body, where, turning, it winds cepha- lad, a little to the left of the median plane, in ascending transverse coils; the terminal portion passes between the testes and thence to the left and dorsally over the acetabulum ; finally it turns to the right and crosses obliquely the distal third of the cirrus pouch (PI. I, Fig. 3), where it enters a well defined metraterm or vagina. The base of the metraterm is enlarged and lies across the left side of the terminal portion of the cirrus pouch, but the neck parallels the pouch and ter- minates anteriorly and to the left of the pouch in the common genital atrium. The wall of the metraterm is thickened and supplied with an inner sheet of circular muscle fibers and an outer sheet of longitudinal fibers. An invagination of the cuticula appears to form the lining of its lumen, the wall of which is transversely ridged. PARASITIC WORMS BERMUDA. I. TREMATODES. 219 A diffuse, but well defined, shell-gland (PI. I, Fig. 6), or gland of Mehlis, lies dorsal, and for the most part anterior, to the ovary. Its posterior portion covers the anterior third of the ovary. The oviduct penetrates the mass of shell-gland cells, which are connected with the oviduct by numerous minute ducts. A globular compact receptaculum seminis (PI. I, Figs. 1, 5, 6), one- third the bulk of the ovary, lies dorsal to the shell-gland and the anterior half of the ovary. A small duct leaves its anterior margin and turning mediad joins the oviduct soon after it leaves the ovary. At the junction of the receptaculum duct with the oviduct a tubular Laurer's canal (Fig. 6) originates and in a slightly undulating course makes its way dorsad and opens on the dorsal surface of the body dorsal to the ovary and the posterior part of the acetabulum and slightly to the right of the median line. The vitellarium (PI. I, Figs. 1, 2, 5) is rather striking in appearance and is composed of two masses of convoluted tubules grouped in moss- like patches, which lie in the lateral and latero-dorsal fields in the middle three-fifths of the body. The patches are fairly definite and constant in number, three patches being present on the right and four on the left side. They extend forward of the anterior margin of the acetabulum a distance approximately half of the diameter of that organ, the posterior limit being about mid-way between the testes and the posterior end of the body. The latero-dorsal patches are dorsal to the testes and uterine coils. A fine vitelline duct (PI. I, Fig. 6) connects with each other the patches or groups of each side and a larger vitelline duct leaves the central group of each side and passes transversely mediad to unite with the one from the opposite side to form a small, but distinct, vitelline reservoir, which lies dorsal to the left margin of the ovary. From the reservoir a small duct leads cephalad and joins the oviduct a short distance beyond the union of the receptaculum duct with the oviduct. The eggs (PI. I, Fig. 4) are numerous, spindle shaped, light brown in color, with thick shell. A comparatively large, well-defined and easily separated operculum is present, and a slight opercular rim can be detected. The opercular pole is the more pointed. The eggs measure 0.0375 mm. to 0.0450 mm. in length by 0.015 mm. to 0.020 mm. in width. The older eggs contain a well developed embryo, but many appear empty, which probably indicates a non-fertile condition. Excretory System. — The excretory system (PI. II, Fig. 7) is volumi- nous and consists of an enormous median dorsal reservoir, with a pair of anterior prolongations. The reservoir is one-fourth the width and 220 BARKER. one-half the depth of the body, and extends from the posterior end of the body to the posterior margin of the ovary, where it bifurcates, one arm passing to the left and one to the right of the ovary; the arms extend cephalad around the oral sucker to the anterior end of the body. The reservoir and arms give off numerous long lateral and deep ventral diverticula, but these do not anastomose. The reservoir terminates behind in a short narrow median canal at the posterior end of the body, which ends in a well-defined excretory pore, terminal in position and nearer the ventral than the dorsal surface. The short excretory canal appears to be lined with cuticula. The entire excre- tory system is filled with a mass of fine globular, gray and golden, glistening particles among which are numerous larger globular bodies which stain a bright blue with methylen blue. 2. Taxonomy. Braun (1901, p. 36) described under the name of Distownvi irrora- iuvi R. a trematode found in the intestine of a sea turtle, ThaJassoch- elys caretta, from New Guinea, which has a number of characters similar to the trematode described in this paper. Looss (1901, p. 558) described a similar trematode found in the stomach of a sea turtle, Thalassochelys corticata, from Triest, which he named Pachypsolus lunatvs. In a later paper Looss (1902, p. 485), after a careful com- parative study of new adult specimens, as well as the forms described by Braun and by himself, reached the conclusion that all were speci- mens of Distowvm irrondum Rudolphi, those described by Braim and by himself in his earlier paper being young forms, while those studied bv himself later were mature. He accordinglv classified all of them as Pachypsolus irroratvs (R.). * Looss (1902, p. 503) gives the following characters for the genus Pachypsolus, "Mittelgrosse Distomen mit sehr kraftigem, dickem, vorn und hinten abgerundetem, auf dem Querschnitte kurz ovalem Korper. Saugnjipfe gross und kriiftig, Haut besonders im Vorder- korper mit scheinbaren Biindelen feiner stabchenartiger Stacheln bewaifnet. Darm mit starkem Pharynx, ganz kurzem Oesophagus und Darmschenkeln, die bis auf einige von ihren Angfangstheilen nach vorn abgehende Blindsacke einfach sind. Excretionsblase Y formig, mit bis zum Keimstock reichendem Stamme und bis ins Kopfende sich erstreckenden Schenkeln. Stamm und Schenkel mit massig zahlreichen, weiten und zum Theil wieder gespaltenen Seitenzweigen, die nach der Bauchseite hinabsteigen mit Ausnahme des vordersten PARASITIC WORMS BERMUDA. I. TREMATODES. | 221 Paares, welches uber clem ]Mun(lsaiignapfe eine einfache Querana- stomose der Schenkel bildet. Genitalporus etwas Hnksseitig von dera Bauchsaiignapfe, Copulationsorgane vorhanden. Cirnisbeutel cy- lindrisch, von betraehtlicher Ltingc, in seinem Innern eine mehrfach gewunden, sehlanke Sanienblase, lange, cylindrische Pars prostatica und dicker Penis, der sich im ausgestiilpten Zustande nach seiner Spitze zu merklich verjilngt. Hodeii stark seitlich hinter dem Bauchsaiignapfe. Keimstock seitlich vor ihnen; Laurer'scher Canal und Receptaculum seminis vorhanden. Dotterstocke in den Seiten und unter der Riickenflache, aus in der Jugend deutlich sternformigen FoUikelgruppen zusammengesetzt, Uterusschlingen hauptsachlich hinter den Hoden die ganze Breite des Korpers ausfiillend und nur die Enden der Darmschenkel freilassend. Eier zahlreich, klein, mit zugespitztem Deckelpol und dickerm Hinterende, zwischen 0,04 und 0,05 mm. lang. Bewohner des Tvlagens von Seeschildkroten. Typus: P. irroratns (R.)." The trematode from ChcJonia imbricata which I have described has, in general, the characters of the genus Pachypsolus, and I do not hesitate to place it in that genus. When compared with the trema- todes described by Braun and Looss under the name Irwratus several essential differences are e\'ident. Externally the following may be noted. The absence of spines, or scales, which may, however, have been lost, the very large and more nearly equal size of the suckers, the ventral cup-like depression and the non-salient genital pore. In- ternally, the position of the testes and ovary nearer the acetabulum and the less diiTuse arrangement of the vitelline masses, which are more nearly like those described by Braun, may be noted. The most striking and essential difference, however, is the size and position of the cirrus pouch, which in Pachypsolus irwratus (PI. II, Fig. 11) bends around the acetabulum, its posterior end extending to the level of, or posterior to, the ovary, while in the form here described (PI. I, Fig. 3) the cirrus pouch is much shorter, parallel with the dorso- ventral axis of the body and entirely anterior to the acetabulum. Linton (1910, p. 24) has described a new species, Pachypsolns omlis, found in large numbers in the intestine of a Loggerhead Turtle {Caretta caretia) from the Tortugas. A third species, Pachypsolus tcrtius, has been described by Pratt (1914, p. 416) from the small intestine of the same host and of the same locality. The species described by Linton and by Pratt differ from P. irroratvs in minor points and distinctively in the position and extent of the cirrus pouch. Pratt (1914, p. 418) describes the cirrus sac in P. tcrtius (PI. II, Fig. 9) "222 J BARKER. as "a long cylindrical structure, extending from the genital pore around the dorsal side of the acetabulum to the vicinity of the ovary and the shell-gland, and in some cases to the anterior border of the testes." According to Linton (1910, p. 25) the cirrus pouch in P. ovalis (PI. II, Fig. 10) is " relatively short, reaching barely to the posterior edge of the acetabulum." Both Linton and Pratt consider the differences in the extent of the cirrus pouch, together A\ath other minor differences, to be of specific rank. It is evident that the form which we have described resembles P. ovalis Linton more than it does P. tertius Pratt or P. irrorahis (R.) Looss; l)ut it differs from P. ovalis Linton in minor characters and distinctively in the position and lesser extent of the cirrus pouch. The difference in the length of the cirrus pouch in P. ovalis Linton and in the trematode here described is greater than that between P. ovalis and P. tertius Pratt and decidedly greater than that between P. tcrtivs Pratt and P. irroratiis (R.) Looss. We agree with Pratt that the "actual position is undoubtedly de- pendent upon the condition of contraction," but it seems improbable that this constant and marked difference could be due entirely to the contraction of the acetabulum or the body. We feel warranted in ascribing to this difference in the position and extent of the cirrus pouch, taken together with the minor differences noted, a specific value, and therefore class this trematode as a new species in the genus Pachi/psolus, designating it as Pachy psoitis hrachus} In the four species of Pachypsolus now recorded we find, in addition to differences of secondary importance, a striking gradation in the position and size of the cirrus pouch, which is the distinctive specific character. The old question, raised by Looss, arises as to specific differences and the specific effects of different hosts on the same species. From the standpoint of geographical distribution, it is of interest to find in the Hawk's-bill Turtle from the Bermudas a different species of Pachypsolus from that found in the Loggerhead Turtles of New Guinea and the Mediterranean and from those found in the Logger- head Turtles of the Tortugas. 1 ..Jpaxi's, short, having reference to the cirrus pouch. PARASITIC WORMS BERMUDA. I. TREMATODES. 223 Synechorchis megas, n. g. et n. sp. (PI. Ill, Figs. 13-22). 1. Morphology. General Appearance. — Twenty-four specimens of this trematode were studied, twelve of which were fixed in 2% formol and twelve in vom Rath's osmio-sublimate mixture. In general the body is boat- or cradle-shaped, the dorsal surface being strongly convex both longi- tudinally and transversely and the ventral surface correspondingly strongly concave. The body tapers slightly toward the anterior end making the posterior end the broader and more bluntly rounded. In unmounted specimens the length varies from 4.2 mm. to 9 mm. and the width from 2.2 mm. to 3.2 mm. The thickness of the body is 1.04 mm. with slight variations. No cuticular spines or scales were found. At the anterior end of the body is a well defined terminal cephalic hood or collar (PI. Ill, Figs. 18, 19), 1.50 mm. to 1.65 mm. wide and 1.05 mm. to 1.17 mm. long. The dorsal margin of the hood is unbroken but the ventral margin is indented by a wide shallow notch or hilus (Fig. 17). The whole hood has the general appearance of a cocked hat; its ventral face is slightly concave, with the lappets not promi- nent, giving the whole somewhat the shape of a kidney. A well defined muscular oral sucker lies in the ventral cupped face of the hood, but an acetabulum is not present (Figs. 17, 18, 22). At the posterior end of the body, on the dorsal surface, in the median line is a funnel-shaped opening, which marks the termination of the excretory system. The male and female genital pores are separate and salient; they lie on the medial side of the left intestinal caecum (Figs. 17, 22) at the level of the posterior margin of the anterior fourth of the body. The large cirrus was extruded in several of the specimens examined (Fig. 17). Digestive System. — The oral sucker, having a fairly well developed musculature, opens directly into the oesophagus. It measured 0.66 mm. wide by 0.60 mm. long. A pharynx is not present. The length of the oesophagus varies much; in some specimens it appears to be wanting, in others it may reach a length of 0.50 mm. The wide intestinal caeca occupy a lateral and dorsal position in the body and extend from the oral sucker in an undulating course to near the end of the body, where they end blindly. The caeca throughout their 224 BARKER. course are folded or pleated, which gives rise to distinct but irregular pockets along their course. Male genitals. — One of the most characteristic features of this tre- matode is the testes (Fig. 22), twelve in number, arranged in two groups of six each. They are small, irregular, lobed bodies situated in the posterior third of the worm lying on each side of the body immedi- ately ventral to the terminal portions of the intestinal caeca. Taken, together the testes have the shape of a horseshoe, with its open end directed cephalad and extending from the level of the ovary and vitelline glands caudad to the ends of the intestinal caeca. The testes may be separated from, or may overlap, one another. A small duct connects all the testes comprising each group and a larger duct, the vas efferens, passes mediad from the anterior testis of each group. The vas efferens from the left side passes transversely across the body and unites with the short vas efferens from the right side. The vasa effer- entia vmiting from a short vas deferens, which passes cephalad in the right mediolateral field and joins a long tubular convoluted seminal vesicle, which runs anteriorly, mediad to the right intestinal caecum, and enters the cirrus pouch (Fig. 22). The seminal vesicle is lined with a high columnar epithelium. The cirrus pouch (Figs. 22, 20) is a very muscular elongated sac lying between the intestinal caeca at the level of the posterior margin of the anterior fourth of the body. It extends obliquely across the body from right-dorsal to left-ventral and enters the male genital pore. It is provided with an outer thick sheet of strong longitudinal muscle fibers and an inner (toward its lumen) thin sheet of circular muscle fibers. Parenchymal tissue fills the space between these muscle sheets. The seminal vesicle enters the base of the cirrus pouch, where it enlarges to form a short tubular pars prostatica, which is surrounded by the prostate cells. The pars prostatica enters a cone-shaped cavity, the ductus ejaculatorius, which is one-fourth of the length of the pouch and is lined with high columnar epithelial cells having the appearance of coarse cilia. The ductus ejaculatoris is followed by a narrow canal which forms the lumen of the cirrus (Fig. 20). The cirrus is strongly developed and consists of two distinct regions, both of which are protrusile. The basal proximal portion is bulbous and in one specimen measured 0.33 mm. long by 0.25 mm. in diameter; the distal portion is more slender and tapering and in the same speci- men measured 0.50 mm. long bv 0.125 mm. in diameter. The distal portion can be retracted into the bulbous portion. The entire extruded cirrus may be 0.85 mm. long. The cirrus is covered with a PARASITIC WORMS BERMUDA. I. TREMATODES. 225 cuticula in which, on the distal portion, are lightly embedded minute spinelets. It is supplied with an outer sheet of circular muscle fibers and an inner thicker sheet of longitudinal muscle fibers. Its lumen is lined with cuticula. The external opening of the cirrus pouch is separate from that of the metraterm, or vagina, and lies mesad to it and to the left intestinal caecum. Female gcniiah. — The OA'ary (Figs. 13, 22) is a little larger than a single testis and is irregular in outline with a lobed margin. It lies at the right of the median plane, posterior to the uterus and at the level of the most anterior testes. A short oviduct leaves the dorso-medial portion of the ovary and, proceeding obliquely dorsad and to the left, is joined by the Laurer's canal, whence it turns posteriad and mediad across the dorsal surface of the shell-gland (gland of Mehlis) and is joined in the central area of the shell-gland, by a duct from the yolk reservoir. The common duct now enters the shell-gland and enlarges, forming the ootype, which receives numerous small ducts froni the shell-gland, after which it passes ventrad and posteriad through the shell-gland to its posterior margin, whence, after making several coils, it turns cephalad along the left side of the shell-gland and con- tinues as the uterus. The uterus makes its way cephalad, in the median area, in wide compact transverse folds, which may extend laterally as far as the outer edge of the intestinal caeca and \'itelline glands. The uterus terminates in a well defined metraterm, or vagina, which opens to the exterior through a separate female genital pore (Figs. 17, 20, 22) at the left of the male genital pore and central to the left intestinal caecum. The metraterm is an elongated slightly convoluted tubular organ, approximately as long as the cirrus pouch, and lies caudad, and almost parallel, to the pouch. Its wall is strik- ingly thick and muscular, being provided with a thick outer layer of longitudinal muscle fibers and a thick inner (toward the lumen) layer of circular fibers. Its lumen is lined with a thick layer of cuticula, which is raised into longitudinal ridges. A compact, irregularly shaped, shell gland, or gland of INIehlis (Figs. 13, 22), as large as the ovary, lies in the median field at the left of, and more dorsal than, the ovary. A receptaculum seminis is not present. A short Laurer's canal lea\'es the oviduct near the ovary and proceeds dorsad and cephalad opening on the dorsal surface at the right of the median line and slightly anterior to the shell-gland. The ^■itellarium (Fig. 22) is composed of two groups of vitelline glands lying in the lateral fields of the third fourth of the body, ventral to the intestinal caeca. Each group is made up of from seven to ten 226 BARKER. compact coarsely granular glands, which are so arranged as to simu- late an anterior prolongation of the free ends of the testicular horse- shoe; they extend cephalad to approximately the same level on the two sides of the body. A small Aitelline duct connects the successive glands of each group ; a larger one leaves the posterior gland of either side and, passing caudad and mediad, unites with its mate to form a yolk reservoir (Fig. 13), which lies dorsal to the anterior portion of the shell gland. A small yolk duct leads from this reservoir and joins the oviduct in the central area of the shell gland. Excretory System. — Two lateral canals, one on each side of the body, arise at the level of the oesophagus. They parallel the sides of the body and lie slightly external and ventral to the intestinal caeca. At the level of the anterior testes their course becomes obliquely caudo- mediad and they unite just posterior to the shell-gland to form the excretory bladder, which lies in the median plane at about the level of the more posterior testes and ventral to the intestinal caeca. The bladder extends backward in a straight course from the shell gland and terminates in the excretory funnel near the posterior end of the body. The funnel itself runs dorso-caudad and opens through an excretory pore on the dorsal surface in the median line 0.15 to 0.30 mm. from the posterior margin of the body. According to Looss (1902 : 593) this excretory funnel (Figs. 14-17, 22) is characteristic of the Prono- cephalidae; it is lined by cuticula raised into 7 to 9 longitudinal ridges. Cilia were not observed in the funnel, the inner end of which is sur- rounded by numerous gland cells. The uterus is packed with numberless eggs; those from different parts of the uterus were studied. In mass the eggs appear dark brown, but individual eggs are light brown or golden yellow. In shape (Fig. 21) they vary from short or long oval to ovoid, and every egg bears a tuft of filaments at each pole. The body of the egg in glycerine preparations varies from 0.0287 to 0.0387 mm. in length, and from 0.0162 to 0.0187 mm. in width. The shell is thick and has at the more pointed end a definite flattened operculum, but mthout an opercular rim. Six to ten coarse filaments, which may attain a length of 7 times that of the egg proper, occur at the opercular pole and there are at the opposite end 12 to 20 coarse very long filaments, 15 times as long as the egg proper, wath an equal number of short finer filaments. The diameter of the coarser filaments is about one-half the thickness of the egg shell and they appear to be composed of the same material. The intertwining of these filaments causes a characteristic massing of the eggs, and makes it difficult to separate them. PARASITIC WORMS BERMUDA. I. TREMATODES. 227 2. Taxonomy. The presence of the single sucker, the cephalic hood, and the peculiar funnel-shaped depression which is associated with the excretory pore undoubtedly place this trematode in the Family Pronocephalidae as characterized by Looss (1902, p. 611). It cannot, however, be placed in the genus Charaxicephalus of Looss on account of the difference in the number, arrangement and position of the testes and other, though minor, differences. It has many of the characters given by Braun (1901, p. 48) for Mo7iostomwn pandum, which he describes as follows: "Mir liegt nur ein einziges, wohl erhaltenes Exemplar vor, das folgende Verhaltnisse aufweist: es ist 11 mm. lang, kahnformig gekriimmt, verbal tnismassig platt, der Riicken gewolbt, die Bauchflache konkav; weder der Hinterrand noch die SeitenrJinder sind wie bei Mon. trigonocephalum bauchwarts eingebogen: am Hinterrande keine Spur von irgend welchen Anhangen. Das Kopfende tragt ein nierenformiges, dem Halskragen der Echinostomen ahnliches Schild (2 mm. breit), aus dem sich ein niedriger, an der Spitze die Mundoflfnung tragender Kegel erhebt: offenbar entspricht dieses Schild dem Kopfwulst der bisher besprochenen Monostomen aus Seeschildkroten, der demnach auf der Ventralflache nur schwach gebogen und nicht winklig ausgeschnitten ist wie bei Mon. trigonocephalum Rud. Die Breite des Korpers betragt in der Hohe der Genitalpori 2.7, am Beginn der Dotterstocke 3.5, und in der Hohe des Keimstockes 4 mm.; sie nimmt also ganz all- mahlich von vorn nach hinten zu. Der Saugnapf ist 0.625 mm. lang und 0.729 mm. breit; vom Oesoph- agus kann ich etwas bestimmtes nicht angeben, da ich ihn nicht sehe, allem Anschein nach ist er kurz, denn die beiden Darmschenkel sind bei genligend starker Vergrdsserung sowohl auf der Riicken- wie Bauchflache dicht hinter dem Kopf schild bereits erkennbar: sie Ziehen, die Endteile der Geschlechtsgange zwischen sich fassend nach hinten und sind zwischen den Dotterstocken und dem Uterus bis an die Hoden zu verfolgen; ihr weiterer Verlauf ist nicht mit Sicherheit zu erkennen, sie scheinen dorsal iiber den Hoden und der Mittellinie etwas mehr genahert bis an den Hinterrand der Hoden sich zu ers- trecken. Soweit ich sie deutlich erkenne, sind sie nach aussen wie nach innen mit kurzen Blindsiickchen besetzt. Vom Exkretionsapparat sind nur die beiden weiten Sammel-rohren aussen von den Dotterstocken erkennbar. Wie haufig bei Monostomidcn, findet sich auch hier je eine Aus- '228 BARKER. miindungsstelle fiir mannliche und weibliche Organe; dieselben liegen dicht neben einander, hinter der Gabelstelle des Darms auf der linken Seite, die Uterusmiindung seitlich von der Cirrusmiindung. Ganz im Hinterende liegen symmetrisch die beiden grossen (bis 3 mm. langen) vielf ach gelappten Hoden ; sie beriihren sich hinten mit ihren medianen Flachen, A-orn weichen sie auseinander. Vom Leitungsweg bemerkt man reehts die gewundene Vesicula seminalis, die durch einen graden Kanal in den langgestreckten und dickwandigen Cirrusbeutel miindet ; seine Lange betragt iiber 2 mm. In dem von den vorderen Enden der Hoden freigelassenen Raume liegt reehts der vierstrahlige Keimstock, neben und etwas hinter diesem in der ISIittellinie die Schalendriise. Hier beginnt der Uterus, auch fliessen an dieser Stelle die queren Dottergange zusammen. Die Dotterstocke Hegen wie gewohnhch seitHch im Korper und erstrecken sich vom Vorderende der Hoden bis vor die Korpermitte; sie bestehen aus zahh-eichen, eine traubige Gruppierung aufweisenden FoUikeln. Die UterusschHngen breiten sich, c^uere Richtung einhaltend, in dem Raum zwischen den Dotterstocken und vor den Geschlechts- driisen aus; das neben dem Cirrusbeutel liegende Metraterm ist kurz vor seiner Ausmiindung von einer kompakten Driisenmasse umgeben. Die Eier scheinen Polfiiden nicht zu besitzen; sie liegen allerdings so dicht im Uterus, dass sich Filamente den Blicken leicht entziehen konnten, andererseits wiirde aber, wenn Filamente vorhanden waren, kaum eine sehr dichte Lagerung der Eier moglich sein ; Messungen an jungen, sicher der Anhange entbehrenden Eieren aus dem Anfangsteil des Uterus ergaben 0.035 mm. Lange und 0.01 mm. Breite." It is evident that the trematode which I have described differs from M. imndum not only in minor dfetails but more especially in the larger number of testes. Pratt (1914, p. 411) has described a monostome trematode, Wilderia cUiptica, found in the Loggerhead Turtle from the Tortugas, which has many characters in common with both M. pandum and the form here described, but differs from both of them in the absence of a cephalic collar or hood. On the ground of the absence of a collar and the presence of several testes Pratt has created the new genus and new species Wilderia eUiptica. The trematode described in this paper cannot be classed as M. pandum, on account of the several testes, nor as Wilderia elliptica, on account of the presence of a definite cephalic hood or collar. If, as Pratt (1914, p. 416) suggests, Braun's description of the testes PARASITIC WORMS BERMUDA. I. TREMATODES. 229 in M. panel um is incorrect and " they are as a matter of fact made up of successive pairs of distinct organs," then the trematode which I ha\'e described may be identical with Monosfomum pandum. Also, if Pratt is in error regarding " the slightest indication of the collar-like cephalic ridge at the forward end of the body " being absent in Wildcria elliptica, his species is probably identical with the trematode which I ha^'e described and with Monostomuvi pandum. Until these points are determined, it seems advisable to create a new genus and a new species for this trematode, which I accordingly designate as Syne- chorchis mcc/as, making it the type species of a new genus, Syncchorchis,^ designed to include those monostome trematodes which have a con- tinuous cephalic collar, and numerous testes placed laterally in the posterior part of the body. The material, on which these descriptions are based, was collected and sent to me by Prof. E. L. Mark, Director, and Dr. W. J. Crozier, Resident Naturalist, of the Bermuda Biological Station. To both of them I wish to acknowledge my appreciation and indebtedness. To Mr. Hiram O. Studley, one of my students, I desire to express my appreciation for his assistance in making a preliminary study and drawings of the second form described in this paper. ^ Swexvs, continuous line, and opxi-s, testicle. k 230 BARKER. Papers Cited. Braun, M. 1901. Trematoden der Chelonier. Mitth. zool. Mus. Berlin, Bd. 2, 58 pp., 2 Taf. Linton, E. 1910. Helminth Fauna of the Dry Tortugas. II. Trematodes. Carnegie Institution of Washington, Publication No. 133, pp. 11-98, 28 pis. Looss, A. 1901. Notizen zur Helrninthologie Egyptens. IV. Ueber Tre- matoden aus Seesehildkroten der egyptischen Kiisten. Centralbl. f. Bakt., Abt. 1, Bd. 30, pp. 555-569. Looss, A. 1902. Ueber neue und bekannte Trematoden aus Seesehild- kroten. Zool. Jahrb., Abt. f. Syst., Bd. 16, pp. 411- 894, Taf. 21-32. Pratt, H. S. 1914. Trematodes of the Loggerhead Turtle (Caretta caretta) of the Gulf of Mexico. Arch, de Parasitol., tom. 16, pp. 411-427. EXPLANATION OF PLATES. All drawings were made with the aid of a camera lucida except Figures 10, 11 and 12. act. Acetabulum o'typ. Ootype cae. in. Intestinal caeca phx. Pharynx can. exc. Excretory canal po. exc. Excretory pore can. L. Laurer's canal po. gen. Genital pore cir. Cirrus po. gen.' Male genital pore coll. Collar po. gen." Female genital pore cstr. vt. Yolk reservoir poc. V. Ventral cup di. vt. Yolk duct rep. sem. Recaptaculum seminis fil. pol. Polar filament sac. cir. Cirrus pouch gl. cnch. Shell gland sue. or. Oral sucker gl. prost. Prostate gland t€. Testis gl. vt. Vitelline glands Ul. Uterus oa. Ovary va. df. Vas deferens oes. Oesophagus vg. Vagina o'dt. Oviduct vs. exc. Excretory vesicle op. Operculum vsl. sem. Vesicula seminalis 232 BARKER. PLATE I. Pachypsolus brachus. Figure I. Figure 2. Figure 3. Figure 4. Figure 5. Figure 6. Ventral view. X 40. Scale line, at the margin, represents 1 mm. Dorsal view of unmounted specimen. X 40. Scale line 1 mm. Cirrus pouch, vagina, acetabulum, etc. Composite view ob- tained by superposing several successive para-sagittal sections. X80. Eggs with and without lid. X 110. Scale line 0.1 mm. Dorsal view. X 40. Scale line 1 mm. Reconstruction of female genitals, based on frontal and sagittal sections. X 60. Barker. — Bermuda Trematodes. Plaie I. Ml-'-ft^ ^^JTS.oU. Fig. 5 Proc. Amer. Acad. Arts and Sciences. Vol. LVII. 234 BARKER. PLATE II. Figure 7. Reconstruction of excretory system of Pachypsolus brachus based on frontal sections. X 50. Figure 8. Ventral view of unmounted specimen of Pachypsolus brachus. X 30. Figure 9. Cirrus pouch etc. of Pachypsolus tertius, after Pratt 1914, Figure 2. X 64. Figure 10. Cirrus Pouch etc. of Pachypsolus ovalis, after Linton 1910, Figure 7. X 60. Figure 11. Cirrus pouch etc. of Pachypsolus irroratus, after Looss 1902, Figure 169. X 38. Figure 12. Reconstruction of digestive tract of P. brachus based on frontal and sagittal sections. X 50. Barker. — Bermuda Trematodes. Plate II. Proc. Amer. Acad. Arts and Sciences. Vol. LVII. 236 BARKER. PLATE III. Synechorchis megas. Figure 13. Figure 14. Figure 15. Figure 16. Figure 17. Figure 18. Figure 19. Figure 20. Figure 21. Figure 22. Reconstruction of female genitals, dorsal aspect, based on frontal and sagittal sections. X 220. Scale line, below the figure, represents 0.1 mm. Sagittal section of excretory funnel. X 250. Cross section of excretory funnel near surface level. X 250. Cross section of excretory funnel at deeper level. X 250. Ventral view of unmounted specimen. X 23. Ventral view" of cephalic region of unmounted specimen. X 68. Sagittal section of cephalic hood. X 75. Details of male and female copulatory organs. X 90. Scale line 0.5 mm. Egg. X 500. Ventral view of mounted compressed specimen. X 33. Scale line 1 mm. Barker. — Bermuda Trematooes. Plate III. Fig. 17 Proc. 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I am greatly indebted to Dr. Mark, both for having made my stay at Bermuda possible and for having given me the opportunity of examining for hydroid material his miscellaneous collections. The only papers hitherto written on the Bermuda hydroids are by Congdon (1907) and Ritchie (1909). Congdon described 19 species, of which five (Eudendrimn hargitti, Clytiafragilis, Sertularella speciosa, Scrhdaria humilis, and Thyroscyphus intcrmcdius) are new. Several others, described by him as new, have been shown by later writers, notably Nutting and Fraser, to belong to already established species, Ritchie discusses the synonymy of one of the Bermuda campanula- rians, and extends the range of two "Challenger" hydroids from the West Indies to the Bermudas. Fraser's paper (1912) on the hydroids of Beaufort, N. C, is also a valuable aid in the study of the Bermuda hydroids, since 21 species of the latter, or over half of the Bermuda forms, occur also in the Beau- fort region. The strong affiliation of the hydroid fauna of the Ber- mudas with that of the West Indian region, already suggested by Congdon, is still more strikingly demonstrated by the species now reported from Bermuda; in all, 29 species are common to the two regions. The distribution of the individual species found in Bermuda is shown in the following table: — 242 BENNITT. Bimeria humilis Eudendrium hargitti .... Eudendrium ramosum . . . Pennaria tiarella Halecium bermudense . . . Halecium nanum ....... *Halecium tenellum Campanularia niarginata 'Campanularia raridentata 'Clytia bicophora 'Clytia cylindrica Clytia fragilis 'Clytia johnstoni Clytia noliformis 'Obelia geniculata Obelia hyalina Lafoea venusia Hebella calcarata Sertularella speciosa *Sertularella tenella Sertularia brevicyathus . . *Sertularia cornicina Sertularia aestuaria Sertularia stookeyi Sertularia versluysi Thyroscyphus intermedius Aglaophenia cylindrata . . Aglaophenia lophocarpa. . Aglaophenia minuta 'Antennularia pinnata .... 'Lytocarpus clarkei Lytocarpus philippinus. . *Monotheca margaretta . . Plumularia diaphana .... Plumularia corrugata. . . . *Plumularia inermis *Plumularia setacea X X a o ^ X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X o s a X X X X X X X X X X X X X X X X X w X X X X X X X X X X X C8 X X X X X X X X X X X X N rrt'.S go X X X X X X X X X X X X X X X X X X X «^ ^m en = a .2 § li a n 0-10 0-10 0-542 0-10 0-10 0-10 0-235 0-440 0-250 0-10 0-25 0-14 0-100 0-10 0-42 0-68 30-324 0-122 0-1 0-103 0-15 0-8 0-10 0-10 0-30 0-10 21-30 24-1181 0-10 0-100 13-201 0-8 0-10 0-576 0-130 0-10 0-106 _ * The first record of these species from Bermuda is contained in the present paper. HYDROID FAUNA OF THE BERMUDAS, 243 This paper records 37 species, including all the hydroids reported from Bermuda up to the present. Where both trophosome and gono- some have been adequately described elsewhere in readily available papers, as is nearly always the case, I have attempted no taxonomic discussion. References are given to the text and plates of the original description, and also to the standard works dealing with the hydroids of the American shores of the Atlantic, in many of which a more com- plete bibliography may be found. Similar references are given for the species which I have not seen, but which have been reported from Bermuda by Congdon and others. Family BOUGAINVILLIDAE. Genus Bimeria. Bimeria humilis Allman. Allman, 1877, p. 8, pi. 5, figs. 3-4. Congdon, 1907, p. 467, fig. 6. Family PENNARIDAE. Genus Pennaria. Pennaria tiarella McCrady. McCrady, 1857, p. 51. Hargitt, 1900, p. 387, 4 pis. Hargitt, 1901, p. 311, figs. 8, 9. Nutting, 1901, p. 337, fig. 14. Congdon, 1907, p. 464. Fraser, 1912, p. 355, fig. 12. Pennaria tiarella is common on the buoys and reefs about Hamilton Harbor and Great Sound, and on the flats outside. Specimens exam- ined showed as many as 17 filiform tentacles, confirming Congdon's belief that P. symmetrica (Clarke, 1879) of Cuba, which has 18 filiform tentacles, is identical with P. tiarella. Stoloniferous reproduction was here observed for the first time in the family, and, so far as I am aware, for the first time in the whole group of Gymnoblastea. ^Yell-marked stolons extend from the distal ends 244 BENNITT. of the stem and branches (Fig. 1) ; they are considerably larger than the parts from which they arise, anastomose freely, and have the same appearance, even to the exact color, as the normal hydrorhiza of Pen- naria. The free ends of the stolons are somewhat knobbed, and along their course appear broken stumps, precisely like the base of the origi- nal stem. This colony was growing in a horizontal position on the under side of a floating buoy, and the stolons had grown along the bottom of the buoy, there to give rise to new colonies. Figure 1. Pennaria tiarella. Colony showing stolon-formation. X If. Family EUDENDRIDAE. Genus Eudendrium. Eudendrium hargitti Congdon. Congdon, 1907, p. 465, figs. 1-5. Besides being extremely abundant in Hungry Bay, the shallow inlet on the south shore where it was found by Congdon, E. hargitti is gen- erally distributed on buoys, timbers, ledges, and eel-grass all over Hamilton Harbor and Great Sound, just below low-tide mark. Cong- don's specimens were 20-50 mm. high, and had 35-45 tentacles. Spec- HYDROID FAUNA OF THE BERMUDAS. 245 imens in my collection from Fairyland Creek reached a height of 80 mm., and the number of tentacles varied from 35 to 60. The distal hydranths are usually larger than the proximal. Eudendrium ramosum (Linnaeus). Tubniaria ramosa, Linnaeus, 1767, p. 1302. Eudendrium ramosum, Hargitt, 1901, p. 309, figs. 5, 6. Eudendrium ramosum, Nutting, 1901, p. 332, fig. 7. Eudendrium ramosum, Congdon, 1907, p. 464. Eudendrium ramosum, Fraser, 1912, p. 349, fig. 8. A few colonies, 50-75 mm. high, were found on a floating buoy in Hamilton Harbor. Family HALECIDAE. Genus Halecium. Halecium bermudense Congdon. Congdon, 1907, p. 472, figs. 16-20. Fraser, 1912, p. 367, fig. 28. Stechow, 1914, p. 134. Stechow, 1919, p. 33. This is one of the most abundant species in Bermuda, growing on a great variety of structures in almost every locality where hydroids are to be found. My specimens attained a height of 75 mm., Congdon's 25-35 mm. Halecium nanum Alder. Halecium nanum. Alder, 1859, p. 355. Halecium marki, Congdon, 1907, p. 474, figs. 21-23. Halecium nanum, Fraser, 1912, p. 367, fig. 29. Halecium nanum, Stechow, 1914, p. 135. Halecium nanum, Stechow, 1919, p. 36. This minute species was often found on floating Sargassum ; a few colonies were also found with //. bermudense on Pennaria from Cow- Ground Flat. The colonies reached a maximum height of 8 mm.; Congdon's specimens were l|-3 mm. high. Halecium nanum appears to have two modes of growth; the resulting I 246 BENNITT. forms are shown to belong to the same species by the presence of the characteristic female gonosome on both. One form is short and scrubby, the other longer and with a few irregular branches coming off just below the hydrophores. The trophosome of the latter variety agrees so well with what Fraser (1912, p. 368, fig. 30) doubtfully called H. repens Jaderholm, that I believe the two are identical, and that he observed this straggling variety of H. nanum. Halecium tenellum Hincks. Hincks, 1861, p. 252, pi. 6, figs. 1-4. Hincks, 1868, p. 226, pi. 45, fig. 1. Nutting, 1901, p. 357, fig. 52. Fraser, 1912, p. 369, fig. 31. Stechow, 1919, p. 41. A few colonies were found, in all stages of growth, on Sargassum at Somerset Bridge, the hydrorhiza forming an extensive network over an alga. The gonosome, essential for a satisfactory determination of the species, which Fraser failed to find in his Beaufort specimens, was present in the Bermuda material. Family CAIViPANULARIDAE. Genus Campanulaeia, Campanularia marginata (Allman). Obelia marginata, Allman, 1877, p. 9, pi. 6, figs. 1, 2. Campanularia insignis, Congdon, 1907, p. 469, figs. 10, 12. Leptoscyphus insignis, Ritchie, 1909, p. 3. Campanularia marginata, Nutting, 1915, p. 44, pi. 6, figs. 5-7. Campanularia raridentata x\lder. Alder, 1862, p. 315, pi. 14, fig. 5. Fraser, 1912, p. 357, fig. 14. Nutting, 1915, p. 39, pi. 4, fig. 1. A single small colony of two or three individuals was found on float- ing Sargassum. Identification is somewhat doubtful, owing to the absence of the gonosome, but the trophosome agrees in every way with Nutting's description. The ten pointed teeth, the 3-5 annula- HYDROID FAUNA OF THE BERMUDAS. 247 tions at the ends of the pedicel, the tubular hydrotheca, and the con- siderable variation in the height of the pedicel, together seem sufficient to place the Bermuda specimen in this species. Genus Clytia. Clytia bicophora Agassiz. Agassiz, L., 1862, p. 304, pi. 29, figs. 6-9. Nutting, 1901, p. 343, fig. 21. Nutting, 1915, p. 56, pi. 12, figs. 1-3. Fraser and many other writers consider Clytia hicophora identical witli C. johnsioni (Alder). Nutting, with some hesitation, regards it as a separate species, on the basis of the following points: 1) the tenuity of the hydro thecal wall; 2) the smaller size of the hydrotheca; 3) the presence of a simple instead of a complex diaphragm. IMy specimens of C. hicophora, found growing on Pcimaria from Cow-Ground Flat, have hydrothecae which are distinctly smaller than those of C. john- sioni, and show many cases of the collapsed hydrothecal wall. They also have only 12 marginal teeth, and there are annulations in the middle of the pedicels, which are sometimes annulated throughout. None of my specimens of C. johnstoni show these features, and I have found no stages intermediate between the two; this seems sufficient to establish C. bicophora as a separate species. Clytia cylindrica Agassiz. Agassiz, L., 1862, p. 306, pi. 27, figs. 8, 9. Nutting, 1901, p. 342. Fraser, 1912, p. 358, fig. 16. Nutting, 1915, p. 58, pi. 12, figs. 6, 7. A few colonies were found on Sargassum at Agar's Island, and on floating Sargassum oft' the north shore. Clytia fragilis C'ongdon. Congdon, 1907, p. 471, fig. 13. Nutting, 1915, p. 62, pi. 15, fig. 1. A number of colonies, 10-12 mm. high, were found on Sargassum at Somerset Bridge. The gonosome was absent, but the trophosome is quite characteristic in this species. k 248 BENNITT. Clytia johnstoni (Alder), Campanularia johnstoni, Alder, 1857, p. 36. Clytia johnstoni, Hincks, 1868, p. 143, pi. 24, fig. 1. Clytia grayi, Nutting, 1901, p. 344, fig. 23. Clytia johnstoni, Fraser, 1912, p. 358, fig. 17. Clytia grayi, Stechow, 1914, p. 128, fig. 5. Clytia johnstoni, Nutting, 1915, p. 54, pi. 11, fig.s. 1-6. Clytia johnstoni, Stechow, 1919, p. 43. This was one of the commonest species on floating Sargassum. The characteristic annulated gonangia were extremely numerous in the specimens collected. In one colony a stolon twice as long as the pedi- cel extended out from the middle of the pedicel, establishing connec- tion with the substratum. This is the first case of stolon-formation that I have seen in the genus Clytia. There were never less than 14 marginal teeth, and I found no cases of the collapsed hydro thecal wall; these points, with the greater size of the colonies, made them readily distinguishable from C. bicophora. Clytia noliformis (McCrady). Campanularia noliformis, McCrady, 1858, p. 92. Clytia noliformis, Nutting, 1901, p. 343, fig. 22. Clytia simplex, Congdon, 1907, p. 472, figs. 14, 15. Clytia noliformis, Fraser, 1912, p. 359, fig. 19. Clytia noliformis, Nutting, 1915, p. 57, pi. 11, figs. 7-10. Colonies of Clytia noliformis are very numerous on floating Sar- gassum. My specimens showed many intergradations between C. simplex as described by Congdon and C. noliformis as described by Nutting. Genus Obelia. Obelia geniculata (Linnaeus). Sertularia geniculata, Linnaeus, 1758, p. 812. Obelia geniculata, Nutting, 1901, p. 351, fig. 38. Obelia geniculata, Fraser, 1912, p. 362, fig. 23. Obelia geniculata. Nutting, 1915, p. 73, pi. 18, figs. 1-5. A few colonies were found on floating Sargassum. HYDROID FAUNA OF THE BERMUDAS. 249 Obelia hyalina Clarke. Obelia hyalina, Clarke, 1879, p. 241, pi. 4, fig. 21. Obelia hyalina, Congdon, 1907, p. 468, figs. 7-9. Obelia congdoni, Hargitt, 1909, p. 375. Obelia hyalina, Fraser, 1912, p. 363, fig. 24. Obelia hyalina. Nutting, 1915, p. 76, pi. 18, figs. 6, 7. This is one of the hydroids found most often on the floating Sar- gassum, and a number of colonies 2-3 cm. high were found on a fish-car at Agar's Island. There were many cases of stolon-formation from the ends of the branches, and in one case these stolons were thickly inter- twined with similar stolons of Aglaophenia minuta. Obelia hyalina often grows far out on colonies of Sertularia stookeyi no larger than itself. Family LAFOEIDAE. Genus Lafoea. Lafoea venusta Allman. Allman, 1877, p. 11, pi. 6, figs. 3-4. Ritchie, 1909, p. 260. Specimens of Lafoea venusta were dredged by the " Challenger" " off the Bermudas, 30 fathoms." Family HEBELLIDAE. Genus Hebella. Hebella calcarata (A. Agassiz). Lafoea calcarata, A. Agassiz, 1865, p. 122. Lafoea calcarata, Hargitt, 1901, p. 387, fig. 24. Hebella calcarata. Nutting, 1901, p. 353, fig. 56. Lafoea calcarata, Congdon, 1907, p. 467. Hebella calcarata, Fraser, 1912, p. 371, fig. 34. 250 BENNITT. Family SERTULARIDAE. Genus Sertularella, Sertularella speciosa Congdon. Congdon, 1907, p. 476, figs. 24-28. Sertularella tenella Alder. Alder, 1857, p. 23. Hartlaub, 1901, p. 63, Taf. 5, figs. 21-24, Taf. 6, figs. 2, 4, 7, 9, 10. Nutting, 1904, p. 83, pi. 18, figs. 1, 2. A large number of colonies, about 9 mm. high (Nutting's specimens were 12.5 mm. high), were found among branching Bryozoa in a col- lection made in 1903 by Dr. A. W. ^Yeysse. The trophosome agrees in every way with that described by Nutting and by Hartlaub; occa- sional branches are given off at right angles to the stem, and the hydro- thecal walls may be nearly smooth or may have six or seven well- marked annulations. The gonangia are one and a half to two times the length of the hydrothecae. Genus Sertularia. Sertularia brevicyathus (Versluys). Desmoscyphus brevicyathus, Versluj^s, 1899, p. 40, figs. 9, 10. Sertularia brevicyathus, Nutting, 1904, p. 60, pi. 6, figs. 1, 2. Sertularia brevicyathus, Congdon, 1907, p. 481. Numerous colonies of this little Sertularia were found on a gorgo- nian stem and on algae dredged in 1903 at four stations on the Challen- ger Bank, about 15 miles southwest of Bermuda, in 31-70 fathoms. Others were collected on Sargassum near Agar's Island. Sertularia cornicina (McCrady). Dynamena cornicina, McCrady, 1858, p. 102. Sertularia complexa, Nutting, 1901, p. 360, fig. 57. Sertularia cornicina. Nutting, 1901, p. 359, fig. 56. Sertularia cornicina, Nutting, 1904, p. 58, pi. 4, figs. 1-5. Sertularia cornicina, Fraser, 1912, p. 374, fig. 38. HYDROID FAUNA OF THE BERMUDAS. 251 Sertularia cornicina was found on the ledges and on Sargassum at both Agar's Island and Somerset Bridge, also on a gorgonian dredged in 32 fathoms on Challenger Bank. The latter specimens showed the for- mation of unusually long stolons from the tip of the colony back to the hydrorhiza. No sign of the often epizoic Hebella calcarata was seen. Sertularia aestuaria Stechow. Sertularia humilis, Congdon, 1907, p. 479, figs. 29-32. Sertularia aestuaria, Stechow, 1919, p. 157. This very common sertularian frequently formed thick mats over the ledges at about the low-tide mark in practically all the locali- ties visited. The specific name humilis was used in 1879 by Arm- strong (Jour. As. Soc. Bengal, vol. 48, p. 101, tab. 9) for Desmoscyphus humilis of the Indian Ocean, and Stechow has suggested for Congdon's S. humilis the name 8. aestuaria, descriptive of its habitat at tide-level. Sertularia stookeyi Nutting. Nutting, 1904, p. 59, pi. 5, figs. 6, 7. Fraser, 1912, p. 375, fig. 39. A large number of colonies, with gonangia, were taken on floating Sargassum both off the north shore and in Hamilton Harbor. In many cases there was profuse growth of stolons from the extremities. Sertularia versluysi Nutting. Desmoscyphus gracilis, Allman, 1888, p. 71. Desmoscyphus inflatus, Versluys, 1899, p. 42. Sertularia versluysi, Nutting, 1904, p. 53, pi. 1, figs. 4-9. Sertularia versluysi, Congdon, 1907, p. 481. Sertularia versluysi, Fraser, 1912, p. 375, fig. 40. Genus Thyroscyphus. Thyroscyphus intermedius Congdon. Congdon, 1907, p. 482, figs. 33-36. 252 BENNITT. Family PLUMULARIDAE. Genus Aglaophenia. Aglaophenia cylindrata Versluys. Versluys, 1899, p. 49, figs. 19-21. Ritchie, 1909, p. 261. Specimens of Aglaophenia cylindrata were dredged by the "Chal- lenger" " off the Bermudas, 30 fathoms." The species is very similar to A. rhyncocarpa Allman, being separated from it by differences in the corbulae. Aglaophenia lophocarpa Allman. AUman, 1877, p. 41, pi. 24, figs. 1-4. Nutting, 1900, p. 92, pi. 18, figs. 6-8. Several immature colonies, about 25 mm. high, were found on the stem of a large colony of Lytocarpus clarkei, dredged in 32 fathoms on Challenger Bank. The gonosome was absent, but the complex trophosome is sufficient for identification. Aglaophenia minuta Fewkes. Fewkes, 1881, p. 132. Nutting, 1900, p. 96, pi. 31, figs. 1-3. Congdon, 1907, p. 483. Fraser, 1912, p. 378, fig. 43. There is a dense growth of this little plumularian on many pieces of floating Sargassum; specimens have also been found at Agar's Island and among material dredged in 32 fathoms on Challenger Bank. The two nematophores noted by Congdon in the axil of each hydro- cladium are mentioned in Nutting's description, though Congdon must in some way have overlooked this statement. No gonosome was found. Genus Antennularia. Antennularia pinnata Nutting. Nutting, 1900, p. 71, pi. 5, figs. 5, 6. Growing among encrusting Bryozoa on a floating buoy in Hamilton Harbor and reaching a height of 37 mm., were a large number of col- HYDROID FAUNA OF THE BERMUDAS. 253 onies of this hydroid, whose canaHculated coenosarc and unprotected gonangia place it in the genus Anicnnularia. The trophosome agrees with that described by Nutting for A. pinnata, except that I was able to find only one nematophore, instead of two, in the axil of each hydro- cladium, and none at all on the stem, although Nutting states that they are "scattered over the stem."^ There is also considerable dis- parity in size between my specimens and his, but this is not conclusive evidence of specific difference. Some of the colonies are sparsely branched, and the arrangement of the hydrocladiu is invariably alter- nate or subalternate. Figure 2. Antennularia pinnata. female gonangia. X 12. Portion of colony bearing male and The gonangia (Fig. 2) are unprotected, oblong-ovate, coarsely and irregularly annulated, about 20 times as long as the hydrothecae, with strictly terminal apertures, and are borne singly on short processes from the stem opposite the hydrocladia. Both male and female gonangia are found in the same colony. The female blastostyle bears usually a single gonophore, which is situated on one side. The male blastostyle is entirely surrounded by the mass of male reproductive cells. The position of the gonangia, their annulated walls, their 1 Professor Nutting has kindly corroborated my identification of this species and of Plumularia inermis. 254 BENNITT. terminal apertures, and their comparatively large size, make this gono- some, previously undescribed, distinct from that of any other American species of Antennularia. Genus Lytocarpus. Lytocarpus clarkei Nutting, Nutting, 1900, p. 124, pi. 32, figs. 5-7. Large colonies of Lytocarpus clarkei, measuring from 100 to 300 mm. in length, were dredged at five stations on Challenger Bank, in 31-70 fathoms. The gonosome is absent, but the trophosome agrees com- pletely with that described by Nutting. The color of the perisarc in the preserved specimens varied from light brown to deep chocolate- brown. Lytocarpus philippinus (Kirchenpauer). Aglaophenia philippina, Ivirchenpauer, 1872, Pt. 1, p. 45, Taf. 1, 2, Taf. 7, fig. 26. Lytocarpus phiUppinus, Nutting, 1900, p. 122, pi. 31, figs. 4-7. Lytocarpus philippinus, Congdon, 1907, p. 484, fig. 37. Lytocarpus philippinus, Fraser, 1912, p. 379, fig. 45. Lytocarpus philippinus, Stechow, 1919, p. 132. An immature colony, about 25 mm. high, was taken on Sargassum at Somerset Bridge. The gonosome was absent. Genus Monotheca. Monotheca margaretta Nutting. Nutting, 1900, p. 72, pi. 11, figs. 1-3. Fraser, 1912, p. 380, fig. 47. Several colonies in good condition, 6-12 mm. high, were found on floating Sargassum. The gonosome is unknown; the trophosome agrees in detail with that described by Nutting. Genus Plumularia. Plumularia diaphana (Heller). Anisocalyx diaphanus, Heller, 1868, p. 42, tab. 2, fig. 5. Plumularia alternata, Nutting, 1900, p. 62, pi. 4, figs. 1, 2. HYDROID FAUNA OF THE BERMUDAS. 255 Schizotricha tenella, Nutting, 1900, p. 80, pi. 4, figs. 4, 5. Schizotricha tenella, Nutting, 1901, p. 365, fig. 70. Plumularia alternata, Congdon, 1907, p. 484. Plumularia alternata, Fraser, 1912, p. 381, fig. 48. Schizotricha tenella, Fraser, 1912, p. 383, fig. 52. Plumularia diaphana, Bedot, 1914, ]). 89, tab. 5, figs. 14-16. Plumularia diaphana, Stechow, 1919, i). 114. Plumularia diaphana is rather common on floating Sargassura. Branches were observed in a few cases, though the colonies are nearly always unbranched. Stechow noticed that in many colonies the proximal three or four hydrocladia were paired instead of alternate; I find this to be almost universally the case in Bermuda specimens. The gonosome is unknown. Plumularia corrugata Nutting. Nutting, 1900, p. 64, pi. 6, figs. 1-3. A few colonies, 10-12 mm. high, were found on floating Sargassum. The gonosome was absent. The colonies were unbranched, and the stem showed a pair of internal ridges at both the proximal and distal; end of each internode. Plumularia inermis Nutting. Nutting, 1900, p. 62, pi. 5, figs. 1, 2, 2a. Fraser, 1912, p. 382, fig. 50. This delicate hydroid covered thickly a large area of eel-grass in the shallow water of Fairyland Creek; the colonies attained a height of 18 mm. The trophosome agrees with Nutting's description, except that the intermediate internodes are much more numerous than one would infer from his reference to their "occasional appearance," and there are often one or two short intermediate internodes between the proximal hydrotheca and the stem. The hydrocladia rarely bear more than three hydrothecae, and are often prolonged into stolons. The gonosome, heretofore unknown, was found in abundance. The gonangia (Figs. 3, 4) are 20-30 times as long as the hydrothecae, unpro- tected, oblong-ovate, decidedly annulated throughout, and differing from those of all other American species of Plumularia in springing directly from the hydrorhiza. The colonies are dioecious; the female 256 BENNITT. blastostyle (Fig. 3) shows the thick, rounded "Deekenplatte" of ectodermal cells about the terminal orifice, and bears usually a single gonophore on one side; the male blastostyle (Fig. 4) is surrounded by a solid mass of sperm-producing cells. Figure 3. Phimularia inermis. Female gonangium. X 11. Figure 4. Phimularia inermis. Male gonangium and portion of colony, showing stolon-formation. X 11. Plumularia setacea (Ellis). Corallina setacea, Ellis, 1755, p. 19, pi. 11. Plumularia setacea, Nutting, 1900, p. 56, pi. 1, figs. 1-4. Several colonies of Plumularia setacea were found on Sargassum; in one group of colonies stolon-formation was extensive. A k HYDROID FAUNA OF THE BERMUDAS. 257 BIBLIOGRAPHY. Agassiz, A. 1865. North American Acalephae. Illustr. Cat. Mus. Comp. Zool. Harvard Coll., no. 2. viii + 234 p. Agassiz, L. 1862. Contributions to the natural history of the United States, vol. 4, 372 p., 35 pis. Alder, J. 1857. A catalogue of the zoophytes of Northumberland and Durham. Trans. Tyneside Nat. Field Club, vol. 3, pp. 1-70. 1859. Description of three new sertularian zoophytes. Ann. and Mag. Nat. Hist., (3), vol. 3, pp. 353-355. 1862. Description of some rare zoophytes found on the coast of Northumberland. Ann. and Mag. Nat. Hist. ,.(3), vol. 9, pp. 311-316. Allman, G. J. 1871. A monograph of the gymnoblastic or tubularian hydroids. Ray Society, London, vol. I, xxiv + 450 p. ; vol. II, 23 pis. 1877. Report of the Hydroida collected during the exploration of the Gulf Stream by L. F. de Pourtales. Mem. Mus. Comp. Zool. Harvard Coll., vol. 5, no. 2, 66 p., 34 pis. 1888. Report on the Hydroida dredged by H.M.S. " Challenger" during the years 1873-1876. Part II. Report Chal- lenger Exped., Zool., vol. 23 (pt. 70), 90 p., 39 pis., 1 map. Bedot, M. 1914. Nouvelles notes sur les hydroides de Roscoff. Arch. Zool. Exper., tom. 54, pp. 79-98, tab. 5. Clarke, S. F. 1879. Report on the Hydroida collected during the exploration of the Gulf Stream and the Gulf of Mexico by Alexander Agassiz, 1877-1878. Bull. Mus. Comp. Zool. Harvard Coll., vol. 5, pp. 239-252, 5 pis. Congdon, E. D. 1907. The hydroids of Bermuda. Proc. Amer. Acad. Arts and Sci., vol. 42, no. 18, pp. 463-485. Ellis, J. 1755. An essay towards a natural history of the corallines. London. 103 p., 40 pis. 258 BENNITT. Fewkes, J. W. 1881. Report on the Acalephae, etc. Bull. Mus. Comp. Zool. Harvard Coll., vol. 8, pp. 127-140, 4 pis. Fraser, CM. 1912. Some hydroids of Beaufort, N. C. Bull. U. S. Bur. Fish., vol. 30, pp. 337-387. Hargitt, C. W. 1900. A contribution to the natural history and development of Pennaria tiarella. Amer. Nat., vol. 34 (401), pp. 387-406. 1901. The hydromedusae. Amer. Nat., vol. 35, (412), pp. 301- 315; (413), pp. 379-395; (415), pp. 575-595. 1909. New and little-known hydroids of Woods Hole. Biol. Bull., vol. 17, (6), pp. 369-385. Hartlaub, C. 1901. Revision der Sertularella-Arten. x\bhandl. a. d. Geb. Naturwiss., Bd. 16, 143 p., 6 Taf. Heller, K. 1868. Die Zoophyten und Echinodermen des adriatischen Meeres. Wien. 88 p., 3 Taf. Hincks, T. 1861. A catalogue of the zoophytes of South Devon and South Cornwall. Ann. and Mag. Nat. Hist., (3) vol. 8, pp. 152- 161, 251-262, 290-297. 1868. A history of the British hydroid zoophytes. London, vol. I, Ixviii -j- 338 p.; vol. H, 67 pis. Jaderholm, E. 1907. Leber einige nordische Hydroiden. Zool. Anz., Bd. 32, pp. 371-376. Kirchenpauer, G. H. 1872. Ueber die Hydroidenfamilie Plumularidae. Abhandl. a. d. Geb. Naturwiss., Bd. 6, 52 p., 8 Taf. Linnaeus, C. 1758. Systema naturae, 10th ed. Holmiae. vol. I, 1384 p. 1767. Systema naturae, 12th ed. Holmiae. vol. I, pt. 2, 1327 p. McCrady, J. 1858. Gymnophthalmata of Charleston Harbor. Proc. Elliott Soc. Nat. Hist., vol. 1, pp. 1-119, pis. 8-12. Nutting, C. C. 1900. American hydroids. Part I. Plumularidae. Spec. Bull. U. S. Nat. Mus., 135 p., 34 pis. 1901. Hydroids of the Woods Hole region. Bull. U. S. Bur. Fish., vol. 19, pp. 325-386. HYDROID FAUNA OF THE BERMUDAS. 259 1904. American hydroids. Part II. Sertularidae. Spec. Bull. U. S. Nat. ilus., 151 p., 41 pis. 1915. American hydroids. Part III. Campanularidae and Bonneviellidae. Spec. Bull. U. S. Nat. Mus., 118 p., 27 pis. Ritchie, J. 1909. Two unrecorded " Challenger " hydroids from the Ber- mudas, with a note on the synonymy of Campanularia insignis. Zoologist, (4), vol. 13, pp. 260-263. Stechow, E. 1914. Zur Kenntnis neuer oder seltener Hydroidpolypen, meist Campanulariden, aus Amerika und Norwegen. Zool. Anz., Bd. 45, pp. 120-136. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas, und anderer Gebiete. Zool. Jahrb., Abt. f. Syst., Bd. 42, pp. 1-172. Versluys, J., Jr. 1899. Hydraires calyptoblastiques recuellis dans la mer des Antilles pendant I'une des croisieres accomplies par le comte R. de Dalmas sur son yacht " Chazalie." Mem. Soc. 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Bennitt, Rudolf. — Additions to the Hydroid Fauna of the Bermudas, pp. 239-259. May, 1922. $.65. 11. Brues, Charles T. — Some Hymenopterous Parasites of Lignicolous Itonididae. pp, 261- 288. 2 pis. May, 1922. $.85. Proceedings of the American Academy of Arts and Sciences. Vol. 57. No. 11. — M.\y, 1922. SOME HYMENOPTEROUS PARASITES OF LIGNICOLOUS ITONIDID^. By Charles T. Brues. SOME HYMEXOPTEROUS PARASITES OF LIGNICOLOUS ITOXIDII)^. 1 By Charles T. Brues. Received Jan. 18, 1922. Presented Jan. 11, 1922. TiiE interesting material which led to the preparation of the present paper was obtained by Professor W. M. Wheeler during the summer of 1920, when he visited the Tropical Research Station maintained by the New York Zoological Society under the direction of Mr. William Beebe at Kartabo, British Guiana. While exploring the forest in the vicinity of the laboratory he found upon the surface of some freshly cut stumps of trees, numbers of a minute species of gall-midge, the females of which were ovipositing in the lumen of exposed vessels of the wood. The larvse of these midges undoubtedly feed within the vessels and their presence attracts swarms of very small Hymenopterous parasites of the family Platygastridse, which are seen scattered over the moist, freshly cut surface depositing their eggs within the bodies of the midge lar^•ui upon which they are parasitic. Professor Wheeler secured speci- mens of the midges and a large series of the parasites which he very kindly turned over to me, thinking that they would prove of interest on account of the extremely long abdomen possessed by some of the parasites, whereby they are enabled to deposit their eggs in the host larvae within the vessels, well below the surface of the wood. The midges are similarly modified for this purpose as the apical abdominal segments are very slender and form an extrusible tubular ovipositor which can also be inserted into the interstices of the woody tissue. Dr. E. P. Felt has been so good as to examine the midges and informs me that they are probably referable to the genus Janetiella Kieffer, although a knowledge of the male might show them to represent a new genus. Felt ('18) lists a number of North American .species of Jane- tiella that produce galls on very diverse plants (Pinaceae, Vitacese and Myricacese) and one that occurs under decaying bark of chestnut, but cites no zoophagous forms. Janetiella occurs in Europe and both North and South America. The parasites proved to be of much greater interest than had been 1 Contribution from the Entomological Laboratory of the Bussey Institu- tion, Harvard University, No. 196. 264 BKUES. anticipated, due not only to theii- strangely modified egg-laying apparatus but on account of a type of variation which they exhibit that appears to be c^uite different from any hitherto reported among the females of insects. Previous to a careful examination it appeared probable that two species were represented in the series, one with the abdomen lengthened to a varying degree in different individuals and a second with the abdomen short like that of most genera of Platygastridse. Closer study has shown, however, that no less than six species, distributed in as many genera, are included, in addition to some males which I cannot associate with the females of any of the species in the lot. These forms from Kartabo, described on a later page, are as follows : Polymecus (Dolichotrypes) minor sp. nov. Synopeas meridionalis sp. nov. Gastrotrypes spatulatus gen. et sp. nov. Polygnotus simplex sp. nov. Platygaster tubulosa sp. nov. Isostasius crassus sp. nov. Three of them, Dolichotriy^pes, Gastrotrypes and Platygaster have the apical portion of the abdomen, the ovipositor, or both, lengthened and modified to reach their hosts within the wood, while the others are apparently in no way specially adapted to the habits of the host. The latter must, therefore, be able to parasitize only host larvse which are feeding very close to the surface of the wood unless they may have active first-stage larvje, but this does not seem probable, since no planidium forms or other free-living larvse are known to occur within the family. ^ In 1911 Crawford and Bradley ('11) described the genus Dolicho- trypes in which they placed a single North American species, D. hopkinsi. This remarkable insect had been first collected in West Virginia by Dr. Hopkins who found the females supposedly o\'ipositing in the bodies of dipterous larvfe living in a stump. Later, in 1897, Professor J. H. Comstock found the same species near Ithaca, New York. He noted them in large numbers, the " females busily inserting the long part of the abdomen into the intercellular spaces of the wood near the bark. They were confined to the outer two inches of the 1 The larvse of some other species of Polygnotus, Platygaster and Synopeas which have been observed are cyclopoid when first hatched, cf. Marchal, '04, Marchal '06, Richardson '14. HITVIENOPTEROUS PARASITES. 265 wood." That the insect occurs generally about Ithaca is evident as it was again taken by Prof. C. R. Crosby in 1910. INIy own first acquaintance with Dolichotrypes was during the summer of 1911 when ]Mr. W. F. Fiske, then director of the Gipsy-moth Laborator^^ at IMelrose, j\Iass., gave me several vials containing numerous specimens of minute Platygastridse that he had collected upon freshly cut stumps not far from his laboratory. He was quite certain that the females were ovipositing in objects concealed within the wood, and from the known habits of numerous related genera, surmised that Dolichotrypes attacks the larva? of some Itonidid midge. ^ At the time I noticed that there was a great Variation in the size and appearance of jNIr. Fiske's specimens and on having them mounted, found that more than one species was represented. This material received no further study, however, till Professor Wheeler showed me the series of similar insects obtained at Kartabo. A re-examination of my New England material then showed the presence of not only Dolichotrypes, but also of two of the other genera found at Kartabo. ]\Ir. Fiske's material then, includes the following: Polymecus (Dolichotrypes) hopkinsi Bradley & Crawford. Synopeas sp. (perhaps S. cornicola Ashm.). Gastrotrypes caudatus sp. nov. I found no males, and there appears to be considerable doubt also, whether the males of Dolichotrypes described and figured by Bradley and Crawford ('11) are really such. This is a minor matter, however, in the present connection, as the interesting points here dealt with relate to the females. If one examines a large series of Dolichotrypes minor under the micro- scope it is at once evident that the individuals vary greatly in length, and that this variation is confined almost entirely to the apical seg- ments of the abdomen. The head, thorax and the first three abdominal segments which form the gaster are uniform in conformation and equal in size, apart from the small differences which are always exhibited even by tlie least variable insects. Among the Platygastridie in particular, \-ariations in body size are usually well marked as the spe- cies are parasites of the larvae of Diptera, and they reflect not only the intraspecific variability of the host, but also to some extent, its change 1 Since then the genus Dohchotrypes has been found in Australia by Dodd who described ('16) a species from Queensland. The single female known was taken on foliage of sugar-cane. 266 BRUES. in size due to age at the time of parasitization. Such variation is always more pronounced among the parasites of insect larvte than is the case with egg-parasites where the food supply of each individual parasite is more evenly apportioned {e.g., in the related family Scelionidse). As can be seen from the diagrammatic sketches shown in Figure 1, the variability in size of the gaster is quite noticeable, but notexcessive. The following segments (four to six) show enormous differences not Figure 1. Polymecus (DoHchotrypes) minor sp. nov. Diagrammatic views in profile of the abdomen of a selected series of females. only in actual length but also in proportionate length in any selected series. Of these, segments four, five and six are black and heavily chitinized, while the seventh or apical one is membranous and almost hyaline. Beyond it, extends the ovipositor and its two sheaths which may be exserted to great length or very nearly concealed within the body. Taken together, these elongated segments resemble the tele- scopic arrangement of parts seen in many insects and other animals. In fact the abdomen of all insects is built upon this principle as its HYMEXOPTEROUS PARASITES. 267 extension and retraction depends upon slight telescopic movements of the sclerites permitted by the infolded membranes which connect their adjacent edges. In a greatly exaggerated form this type of construc- tion is by no means rare; it occurs in the apical part of the abdomen of the higher Diptera, in the midge upon which Dolichotrypes is parasitic, in the Serphoid Scorpioteleia, referred to on a later page, and quite frequently in association with the ovipositor of various insects. Careful dissections of the abdomen of Dolichotrypes show, however, that only the membranous apical segment is extrusible and retractile in response to muscular impulsion. The basal tubular segments (4, 5 and 6) are of a fixed length in each individual insect although one seg- ment may be eight or ten times as long in one specimen as the corre- sponding one in another example. Furthermore, it is impossible to segregate a large series of indi\dduals into classes, based on length of segments as the proportionate lengths are not constant, although there is a well-marked tendency for all to be either long, short, of medium length, etc. The reasons for believing that the lengths of the chitinized segments cannot be changed by muscular action are very clear. The individuals do not show any segments in which the chitinized basal end is tele- scoped within the apex of the preceding segment, nor do any of them show an elongation of the intersegmental membranes. In all cases the exposed portions are black and thickly chitinized, but no hardened portions remain concealed. It is evident, therefore, that the seg- mental lengths of adults are fixed and that they have been determined previous to the hardening of the exoskeleton which occurs soon after the insects have undergone their last ecdysis from pupa to imago. AVhether it occurs at the time of pupation cannot be stated definitely as no puppe have been observed, but as the form of such parts is usually determined at that time there is no reason to believe otherwise in this case. It seems probable, therefore, that the ultimate form of the abdomen is determined when the pupa is first formed, after which pigmentation and chitinization develop slowly. It is noticeable in specimens with extremely long fourth segment that this segment is just long enough, if it could be retracted within the body, to reach to the anterior region of the thorax as is the case with the ovipositor in Inostemma. ^ Such is also true in most of the long-tailed individuals with regard to the length of the membranous scA'enth segment which when exserted equals approximately the sum 1 The condition of the ovipositor in this genus is discussed on a later page (p. 280). 268 BRUES. of the lengths of the more anterior parts of the abdomen and the thorax. In other genera {e.g., Scorpioteleia, p. 279) with similar parasitic habits, the apical portion of the abdomen consists of long tubular slender segments which telescope one within another, but remain movable during life. In Dolichotrypes a precocious extrusion of the segments at the time of pupation would lead to their chitinization and fixation at whatever length they happened to have been protruded. I am therefore inclined to believe that the polymorphic conformation of the abdomen of the imago is actually determined by the individual insect at the time it pupates and that the process is by no means an entirely passive one. Nevertheless, the condition of the abdomen in Dolichotrypes recalls the high and low males of other insects {e.g., certain Dermaptera and lamellicorn beetles) well known to entomologists and subjected to statistical study by Bateson, '92). That this dimorphism may be due to Sporozoan parasites was suggested by Giard ('94), but however plausible and attractive this hypothesis may appear, it seems, at least in the case of the earwigs, to be disproved by the findings of Brindley and Potts ('10) and of Brindley ('18) as these authors found no such correlation between gregarine parasites and the high and low males of Forficula auricularia. So far as Dolichotrypes is concerned such an explanation undoubtedly cannot apply. I have been unable to find any Protozoan or bacterial parasites in them and, moreover, as such endoparasitic species do not have extensive opportunity to acquire microorganisms they are never generally supplied with them, and stand in marked contrast to the free living earwigs, termites, lamellicorn beetles, et al. As already indicated, one of the species of Dolichotr^^jes and most of those belonging to the other genera are new to science so that it has been necessary to include a taxonomic account of these. This is given below. Polymecus (Dolichotrypes) minor sp. nov. 9 . Length 0.8 mm., exclusive of the 4th, 5th and 6th abdominal segments; these together fully exserted 3 mm., and fully retracted 0.4 mm.; the hyaline 7th segment extrusible to 1.5 mm., rarely to 2.5 mm., filaments of ovipositor extrusible to 2 mm., rarely a little more. Black, with the basal half of the scape, the coxae, and the legs, except the thickened parts of the tibiae and femora, brownish yellow. Wings entirely hyaline. Head, oval, fully twice as wide as thick, the occiput more convex than the front; ocelli in a curved line, the lateral ones one-half as far from the eye-margin as from the median one. Head HYMENOPTEROUS PARASITES. 269 shagreened, more densely so above. Eyes bare; malar space half as long as the eye, without furrow. Antennse 10-jointed; scape half as long as the pedicel and flagellum together, much thickened apically; pedicel narrow at base, twice as long as thick; four funicle joints much more slender than the pedicel, the first and fourth short, quadrate, and the second and third considerably longer than thick; club 4- jointed, joints of about equal length, as broad as long. Mesonotum shining, very delicately shagreened, distinctly longer than wide; parapsidal furrows obsolete, indicated only by a depressed spot on the hind margin of the mesonotum; basal scutellar groove narrow, but deep. Scutellum highly convex, with a short, slightly curved and upturned thorn at apex. Pro- and mesopleurse smooth and shining; metapleura punctate and densely hairy, as is also the first abdominal segment; lateral angle of propodeum with a long, straight, backwardly and outwardly directed slender spine. Second segment of abdomen almost as broad and as long as the thorax, polished, with scattered, short, white bristles; broadest just before the middle; third minutely punctured, narrowed apically, the tip only one-fourth as wide as the base of the second. In fully extruded specimens the following seg- ments are very slender, and proportioned as follows, fourth as long as the remainder of the body, the fifth and sixth each as long as the body, including the third segment. In retracted specimens the fourth seg- ment may be only half as long as the second and the fifth and sixth each not over one-half to two-thirds as long as the second. The ovi- positor is rarely extruded to any extent, except in otherwise greatly extended examples. Fourth to sixth segments shining, but under high magnification, distinctly scabrous ; on these segments the sharp lateral edge is visible, but becomes obsolete on the second, except at the extreme base. Fore wing with only very minute marginal cilia; disc hairy, the hairs large and sparse, forming indistinct lines; basal third with minute hairs. Hind wing with two frenulum hooks. Type and numerous paratypes from Kartabo, British Guiana, August 20 and 21, 1920, ovipositing as previously described in a cvit stump, containing larvae of the Itonidid, Janetiella sp. This species differs from D. hopkinsi Crawford and Bradley- ('11) by its much smaller size, almost entirely obsolete parapsidal furrows and somewhat different color. The club of the antenna and the fifth and sixth segments ^ of the abdomen are entirely black, not brown as in the North American species. 1 Not the fourth and fifth as stated by Crawford and Bradley; the third is short and narrow and so closely attached to the second that tliej' have con- sidered it as a part of the latter. 270 BRUES. Polymecus (Dolichotrypes) hopkinsi Crawford & Bradlev ('11, p. 124). Mr. W. F. Fiske obtained numerous females of this species on May 19 near Boston, on cut stumps, behaving as described by Crawford and Bradley. From some of his specimens which he kindly gave me at the time, I have been able to compare the South and the North American species. The genus Dolichotrypes is probably not distinct from Polymecus according to Mr. Fonts who has given much time to taxonomic studies in this family. I have retained it above in subgeneric form to include the two species here dealt with. Other species of Polymecus have the apical prolongation of the abdomen to a lesser degree as do also some species in other genera such as Sactogaster. In addition to the " tail," the females of the latter genus possesses a sac-like enlargement of the venter probably associated with the egg-laying apparatus. A Euro- pean species was bred more than half a century ago by Winnertz ('53) from Contarinia (Cecidomyia) pisi, but further observations on this interesting genus do not appear to have been made. In two other Serphoid families there is a somewhat similar ventral swelling of the second segment which extends forwards ; the Diapriid Cardiopria Dodd and the Belytids Acanosema Kieffer and Cardio- spilus Kieffer are thus modified. Gastrotrypes, gen, nov. Antennre 9-jointed, with a minute hyaline joint-like connection in addition, between the pedicel and first flagellar joint. Maxillary palpi consisting of two equal, elongate joints; labial palpi one-jointed, elon- gate. Head about twice as broad as thick, a little wider than the thorax. Ocelli in a broad triangle, the lateral ones much closer to the eye than to the median ocellus. Parapsidal furrows wanting. Scutel- lum highly convex, without spine, not deeply separated from the mesonotum which bears a large shallow impression at each side behind ; pubescent, especially at the sides apically. Abdomen with the first four segments forming an oval mass; second segment quadrate; third and fourth short, much narrowed; fifth very narrow, sometimes very much elongated; sixth segment membranous, very slender, capable of being greatly extruded. Wings veinless, with very weak discal hairs; with prominent marginal cilia apically behind. Type species: G. spatulatus sp. nov.; other included species: G. caiidains sp. nov. HYMENOPTEROUS PARASITES. 271 Gastrotrypes spatulatus sp. nov. 9 . Length 0.8-1.20 mm., exclusive of the very slender hyaline apical portion of the abdomen which may be entirely withdrawn or extruded to a length of slightly more than that of the body; true ovi- positor very short, never extruded for more than a very short distance beyond the hyaline tube which comprises the sixth and seventh seg- ments, although in many specimens it appears to consist of only a single segment, the sixth. Black; antenna! scape honey-yellow, except more or less at apex; legs honey-yellow, hind coxse sometimes darker; hind femora and tibipe infuscated apically, also sometimes the femora and tibire of the other legs. Head, seen from above oval, slightly, but distinctly more than twice as broad as thick. Ocelli large, the lateral ones removed by somewhat less than their own diameter from the eye-margin, surface of head shining, the vertex and sides of the front shagreened, but the front almost entirely smooth medially; head behind and cheeks, smooth; malar space rather long, more than half the width of the eye, smooth and polished, without furrow. Antennae nine-jointed, not taking into account a minute hyaline con- nection between the pedicel and first flagellar joint; scape about half as long as the remaining joints together; pedicel short, one-half longer than wide; first flagellar joint slightly longer than the second, nearly twice as long as thick; third very small, half as long and half as thick as the second; club 4-jointed, rather stout, the joints as long as broad,, except the last which is more elongate. Mesonotum shining, sparsely clothed with pale appressed hairs laterally; without parapsidal fur- rows; much narrowed anteriorly, with the pleurse largely visible from above. Scutellum not separated by a basal groove, but with a large pubescent fovea at each side; strongly convex, the posterior portion obliquely sloping, almost truncate; entire surface densely hairy and apparently closely punctate beneath the hairs. First four abdominal segments smooth, forming an oval mass as long as the thorax; first segment short, campanulate, with a deep fovea at each side, densely pubescent, except above; medially with a median groove which receives a corresponding ridge on the propodeum when the abdomen is bent upwards; second segment as long as wide, broadest behind, one half longer than the third and fourth together; third and fourth sharply narrowed, of equal length; fifth segment longitudinally aciculate, slightly longer than the fourth, only one-fifth as wide as the second segment, its sides parallel, except on the narrowed apical half; from its apex projects the hyaline sixth segment which is scarcely 272 BRUES. thicker than the posterior tarsi. Pleuree smooth, the metapleurse behind densely clothed with backwardly directed pale hairs. Wings hyaline, with a well-developed marginal fringe apically below; disc with very minute hairs ; hind wing with two frenulum hooks. Type and numerous paratypes from Kartabo, British Guiana (W. M. Wheeler). The color of the legs in this species varies as in Synopeas meridionalis, but the variation is continuous and no color forms are distinguishable. Gastrotrypes caudatus sp. nov. 9 . Length 2.5 mm., including the long stylate apical segment of the abdomen which is nearly as long as the remainder of the body. Black, the wings hyaline, the antennse and legs, including coxse, brownish yellow; upperside of antennae, especially' the apex of scape and the club and the thickened parts of the femora infuscated. Head seen from above twice as wide as thick, shining and almost smooth, the occiput faintly transversely striate and separated from the vertex by a distinct, very fine, raised line; lateral ocelli twice as far from the median one as from the eye margin. Antennae 9-jointed, the scape rather slender, more than half the length of the flagellum; pedicel small, narrower and one-third shorter than the first flagellar joint; first flagellar twice as long as thick, considerably longer than the third; fourth minute, narrowed at base, about as long as wide; club 4-jointed, first joint longest, twice as long as thick, following subequal, each shorter and somewhat thicker than the first; malar space nearly as long as the width of the eye. jNIesonotum smooth and shining, with a band of sparse appressed pale hairs on each side of the middle; parap- sidal furrows entirely absent; behind with a transverse impression on each side at the base of the scutellum, densely clothed with pale hairs. Scutellum highly convex at the middle, obliquely sloped behind and finely, densely punctate beneath a mat of woolly hair. Pro- and mesopleurse smooth and highly polished, metapleura with backwardly directed pale hairs. Propodeum angularly produced laterally and medially with a tubercle which corresponds to a central impression on the dorsal surface of the first abdominal segment. Body of abdomen ovate, nearly as long as the head and thorax together, consisting of four segments; first segment about as wide as long, much narrowed basally, its posterior edge set into the base of the second; medially with a quadrate impression and woolly laterally; second segment as long as wide, broadest behind the middle; its basal margin produced HYMENOPTEROUS PARASITES. 273 forward at the sides and also medially, so that it is bisinuate, on each side of the middle basally with about five fine longitudinal strife that extend nearly half way to apex; third and fourth segments evenly narrowed to the base of the stylate fifth; third twice i^s wide as long; fourth triangular, as long as broad; fifth of even widtli throughout, ten or twelve times as long as broad; apical segments membranous, very slender, usually retracted but in one specimen exserted to a length equally the entire length of the body; filaments of ovipositor not much extruded. Fore wings without marginal cilia; hind wing with two frenulum hooks. Type and numerous paratypes obtained by Mr. W. F. Fiske on stumps of freshly cut trees in the environs of Boston, Mass., on May 19, 1911. This species is very much larger than the preceding South American G. spatidatus described on a previous page, the fifth abdominal seg- ment is much longer and more slender and the sculpture of the second segment is quite diiferent. The length of the fifth segment gives it quite a difl^erent appearance, but as most of the important structural characters are similar in the two forms, I believe that they should be considered to be congeneric. This species was found associated with Dolichotrypes by Mr. Fiske, but does not seem to have been represented in the material examined bv Crawford and Bradlev when thev described the latter ('11). Synopeas meridionalis sp. nov. 9 . Length 0.7-0.8 mm.; occasional specimens from 0.6-0.9 mm. Black; with the basal half of scape, trochanters base of femora and tibise deep yellow; front and middle tarsi whitish, posterior ones in- fuscated; thickened apical parts of femora and tibiae piceous; coxse piceous, the hind ones somewhat lighter; wings hyaline. Head, seen from above, broadly oval, twice as broad as thick; shagreened above, less distinctly so and more shining on the front; lateral ocelli twice as far from the median one as from the eye margin; malar space as long as half the width of the eye; cheeks and back of head distinctly shagreened. Antennae 10-jointed; scape nearly half as long as the remaining joints together, strongly thickened apically; pedicel as long as the width of the scape, nearly twice as long as thick; basal four joints of flagellum slender, the first and fourth but little longer than tliick, second and third longer, but less than twice as long as thick; four club-joints broad, each about as long as thick, oblique. 274 BKUES, Mesothorax as broad as long; the mesonotum strongly convex, shining, faintly shagreened, with only very slight traces of parapsidal furrows anteriorly; scutellum triangular in outline, highly convex and sepa- rated from the mesonotum by a deep, narrow groove, terminated by a short spine or thorn at the apex. This thorn is straight or slightly curved downward at tip and projects horizontally in the plane of the upper surface of the thorax; below it near the upper edge of the propodeum on each side is another backwardly projecting spine of very slender form which is more or less concealed in dried specimens by the dense white backwardly directed hairs that cover the propo- deum; upper surface of thorax sparsely covered by short sparse white oppressed hairs. Pro- and mesopleurje smooth and highly polished, bare. Abdomen short, ovate, no longer than the thorax, highly polished, with a few minute appressed white hairs; lateral carina distinct; first segment densely white woolly pubescent on the sides, but bare medially above; second segment one-third longer than the remaining ones together, widest just before apex, almost as wide as long; remaining segments very rapidly narrower; third and fourth extremely short, fifth noticeably longer; sixth triangular, as long as the three preceding. Ovipositor very short, never exserted for more than one-third the length of the abdomen; third, fourth and fifth segments occasionally exserted to twice the extent described above. Wings with only extremely minute marginal cilia; disc with rather sparse strong hairs which form very irregular lines; base with very minute hairs; anterior and posterior margins bare near the middle of the wing. Frenulum consisting of two hooks. Type and numerous paratypes from Kartabo, British Guiana (W. M. Wheeler). Synopeas meridionalis var. clara var. nov. 9 . Length the same as the typical form. Differs by the entirely yellow antennal scape and yellow coxse and legs, with only the thick- ened part of the hind femora and the hind tarsi, infuscated. Rarely the hind femora are slightly darkened at tips, but there are no inter- grades in the many specimens of both forms before me. The typical form and the pale variety are about equally numerous in the series and the variety averages a trifle larger in size. This species differs from the North American form with similar habits, perhaps S. cornicola Ashm., mentioned on a later page in the following characters: it is considerably smaller, the lateral ocelli are closer to the eye-margin, and the parapsidal furrows are not complete. / HYMENOPTEROUS PARASTIES, 275 Synopeas (?) cornicola Ashm. ('93, p. 288). With Dolichotrypes and Gastrotrypes, Mr. Fiske took several specimens which apparently belong to this species, so far as Ash- mead's original description is concerned. His types of S. cornicola were reared from an Itonidid gall on Cornus iMniculaia, however, which suggests that the present form is probably distinct. These specimens are very much like the species from British Guiana (S. meridionalis) but considerably larger and otherwise specifically dis- tinct. Polygnotus simplex sp. nov. 9 . Length O.S-0.9 mm. Black; antennal scape honey-yellow, darkened toward apex; legs honey-yellow, the coxre piceous and the femora, especially the hind ones inf uscated ; hind tibiee and sometimes the other tibiae also, darkened at tips. Head very much flattened, seen from above nearly three times as broad as long; lateral ocelli nearly as close to the median one as to the eye-margin; head behind the ocelli transversely aciculate; front smooth and polished; malar space long, fully half the width of the eye, faintly shagreened as is also the head behind the eyes. Antennae 10-jointed; scape stout, not much thickened apically, half the length of the remaining joints together; pedicel elongate, nearly twice as long as thick; first flagellar joint small, pale colored; half as long and half as thick as the pedicel; second and third larger, nearly equal, each a little longer than thick; remaining five forming a rather slender club of which the basal joint is smaller and the last longer than the intermediate n;pre or less quad- rate ones. Mesonotum convex, smooth, without furrows, thinly clothed with appressed pale hairs. Scutellum very highly convex, separated at base by a narrow groove, with a large oval impression at each side; its surface finely and shallowly punctate-reticulate. Abdo- men elongate ovate, widest near the apex of the second segment. First segment quadrate, as long as the scutellum, coarsely longi- tudinally fluted; second considerably longer than wide, twice as long as the following segments together, with several short grooves medially at base and a large cuneate basal impression near each side at base; third and fourth equal, each very short; fifth small, triangular, but longer than the fourth; terminal segments not exserted, ovipositor very short. Pro- and mesopleurte smooth and shining; metapleura? clothed with rather sparse backwardly directed pale hairs. Lateral 276 BRUES. carina of abdomen sharply defined. Femora, especially the hind ones, very strongly clavate. Wings with a well developed marginal fringe apically, especially near the lower outer angle; disc with very minute hairs, almost bare, except on apical third of wing where the hairs are large and strong; basal third also somewhat more noticeably furnished with minute hairs. Hind wing with two frenulum hooks. Type and a number of paratvpes from Kartabo, British Guiana (W. M. Wheeler). This species is not very abundantly represented in the collection; a careful sorting has disclosed only about thirty specimens. Platygaster tubulosa sp. nov. 9 . Length to tip of third abdominal segment 1.0-1.2 mm. Black; scape of antenna? yellowish brown basally; coxre, thickened portions of femora and of middle and hind tibia? piceous; trochanters and anterior tibijie pale yellow; tarsi pale, with the last joint infuscated; wings hyaline; third segment of abdomen brown, the apical ones not always extruded, hyaline, with the tips brownish. Head rather flat, more than twice as broad as long when seen from above; ocelli rather large, the lateral ones almost as far from the eye-margin as from the median one; vertex and occiput fineh- shagreened; front shining, practically smooth; malar space nearly half as long as the eye-width, smooth, as is also the head behind the eyes. Antennse apparently 9-jointed, but really with ten joints, counting the small joint just beyond the pedicel; scape considerably more than half as long as the remaining joints together, slender, much narrowed basally; pedicel one half longer than thick; first flagellar joint almost as long as the pedicel, with a short, indistinctly separated ring-joint attached to its base; second and third joints slightly shorter, about quadrate; re- maining four joints forming a slightly thickened, but not very distinct club, the joints of which are slightly longer than wide. Mesonotum with finely impressed, but complete and distinct parapsidal furrows, shining, clothed with sparse appressed hairs. Scutellum large, con- vex, longer than wide and gently sloping downwards behind, its base without foveje and separated only by a thin, shallow impressed line. Propodeum with a strongly raised longitudinal ridge toward each side, enclosing a deep quadrate median impression. Pro- and mesopleurse shining, faintly punctate; metapleura thinly pale pubescent. Abdo- men elongate, lanceolate; first segment broader than long, coarsely longitudinally striated; second segment three times as long as broad HYMENOPTEROUS PARASITES. 277 tapered from middle toward base and apex, more strongly so behind; base medially with some short fine longitudinal striee and toward each side with a larger and deeply impressed groove that extends to the basal fourth; third segment one-third as long as the second, very narrow and evenly contracted to the more or less blunt apex; fourth, fifth and sixth segments tubular capable of being entirely retracted; the fourth nearly as wide as the tip of the third, but the other two very slender; ovipositor never extending much beyond the sixth segment. In the most fully extruded example the fourth to sixth segments together measure 1.5 mm., or distinctly more than the length of the remainder of the abdomen. Wings absolutely hyaline, without dis- tinct marginal fringe, except for some almost transparent hairs api- cally; disc bare. Hind wing with two delicate frenulum hooks. Type and eight paratype specimens from Kartabo, British Guiana (W. M. Wheeler). This species is represented by fewer specimens in the collection than any of the others. Isostasius crassus sp. nov. 9 . Length 1.0 mm. Black; coxae piceous, the trochanters, base of femora and tibiae and tarsi, except last joint pale yellowish; re- mainder of legs dark brown or piceous. Wings hyaline, the vein fuscous. Head considerably broader than the thorax, flattened, more than twice as wide as long when seen from above. Ocelli large, close together, the lateral ones as far from the eye-margin as from the median one; vertex and occiput shagreened, subshining; face shagreened, but more shining; lower margin of face elevated at the insertion of the antennae; makir space indistinctly transversely striated, more than half as long as the width of the nearly round eyes; head behind eyes shagreened, with a faint trace of striae curving upwards across the cheeks. Antennae 10-jointed, less than half as long as the remaining joints together, rather stout, more slender basally; pedicel large, almost as broad as the scape, one-half longer than wide; funicle, con- sisting of the first four flagellar joints, short, the joints very small, of about equal length and quadrate, except for the broader and dis- tinctly transverse fourth joint, club large, first joint narrower than the others; second and third very broad, nearly twice as wide as long, last elongate, triangular, narrower than the preceding one. Thorax broad, the mesonotum as broad as long and the pleurae only slightly visible from above; mesonotum shagreened; rather dull, sparsely clothed 278 BRUES. with fine appressed hairs like the remainder of the thorax and the head; parapsidal furrows deHcate, quite distinct behind, but obsolete in front. Scutellum strongly convex medially, noticeably elevated above the level of the mesonotum, separated at the base by a narrow impres- sion, wider at the sides; posterior margin defined by a semicircular raised margin inside of which is a deep submarginal groove. Propo- deum trilobed behind, woolly on the sides and with a pair of longi- tudinal ridges on its central portion. Abdomen short, ovate, barely longer than the thorax; first segment short, more than twice as broad as long, slightly woolly and longitudinally fluted; second segment shining, very convex, twice as long as v/ide, broadest at the middle, and forming the entire gaster except for the small, elongate-triangular third segment; second finely longitudinally striate at its extreme base, with a larger and deeper groove at each side of the base; third segment punctulate, with sparse pale hairs. Venter highly convex, the lateral carina not very distinct. Propleurse shagreened ; mesopleura broadly impressed medially and behind, obliquely striate near its posterior border; metapleura thinly clothed with short pale hairs. Ovipositor only slightly projecting, curved downwards. Legs rather slender, the femora strongly clavate, the tibiae more weakly so. ^Ying with a short, but distinct marginal fringe, the disc bearing strong and rather large hairs except at the base; vein capitate, one-third as long as the wing. Type and 12 paratype specimens from Kartabo, British Guiana (W. M. Wheeler). As may be gleaned from the taxonomic description (p. 270) of Gastro- trypes, both species have the abdomen lengthened as in Dolichotrypes, but the stylate fifth segment is not of variable length so that, exclusive of the greatly elongated apical membranous segments, all individuals are of approximately equal length. The membranous parts naay be entirely retracted or extruded to a length equalling that of the entire remainder of the abdomen. ^^ Similarly in Platygaster tuhulosa the apical abdominal segments (in this case the fourth and following) are tubular and capable of complete retraction or of extrusion to a length somewhat greater than the re- mainder of the abdomen. The foregoing observations on Dolichotr;v^es and the Gastrotrypes and Platygaster associated with it in British Guiana, suggested an examination of several other Serphoid Hymenoptera. A brief account of these is given below. A similar elongation of the terminal portion of the abdomen occurs in females of members of the genus Serphus (Prodotrypes), but here the HYMENOPTEROUS -PARASITES. 279 anatomical structure is ciuite different. The fifth apparent (possibly really the sixth) segment is contracted to the tip from which issues a stylet-shaped piece, often curved or hooked at the apex and varying in length from a slight projection to a piece nearly as long as the remainder of the abdomen. The stylus is heavily chitinized and appears to be the terminal abdominal segment. Dissection shows, however, that it is composed of the paired sheaths of the ovipositor. These are crescentic in cross-section and fit closely together along the median line above and to form a hollow tube through which the ovipositor extends. The latter can be only slightly extruded as it is enlarged into a bulb at the base which lies within the last segment and is sup- ported by a chitinous strut ventrally. This apparatus is evidently suited for puncturing cjuite resistant tissues. ' In Scorpioteleia of the related family Belytidse an elongation of the terminal abdominal segments occurs, very similar to that sho\)'n by the species of Gastrotrypes and by Platygaster tubulosa. This re- markable genus was first described by Ashmead ('97) from Eastern Canada and later recorded by the present writer ('09) from Wisconsin and the state of Washington. Several other species are known from Europe, which Kieffer ('10) regards as congeneric with Cinetus be- lieving that the modified abdomen of the female is not a good generic character. In the type species, S. mirabilis Ashm., I find upon re- FiGURE 2. Scorpiotelia mirabilis Ashm. Abdomen of female in profile. examination that the apical prolongation of the abdomen is undoubt- edly retractile as it is not chitinized except toward the apices of the segments and the proportionate lengths of the extruded parts of the latter vary considerably in different individuals. The third, fourth, and fifth segments are tubular, successively smaller, but the sixth and last is of much greater diameter, enlarged at the base, then constricted and then turned upward at the pointed tip. Althougli the curve is reversed in position, the resemblance to the sting of a scorpion is very striking and suggested the appropriate name of Scorpioteleia. Dis- section shows the last segment to consist of a ventral valve and two dorsal ones, one overlapping the other. The basal piece extends 280 BRUES. halfway to the tip and the apical one to the tip where it meets the tip of the ventral one. Between these the ovipositor issues. It is very short and its basal attachment lies well within the last segment. Its valves are heavy and lie one on each side, meeting on the median line above and below. Many Belytids are parasitic on dipterous larvse in fungi and quite probably the Scorpiotelei is modified to reach its host in some tube-bearing fungus such as Boletus or Polyporus. ^ In cer- tain other Belytids, e.g., Miota, the tip of the abdomen is upturned and more or less plowshare-shaped, but does not exhibit such excessive elongation. The Platygastrid genus Inostemma is characterized by a most remarkable projection which arises from the dorsum of the first abdominal segment and extends forward over the thorax with its tip over the anterior ocellus. The curvature of this horn corresponds closely to that of the mesonotum, above the surface of which it is raised, and the vertex of the head bears a median depression to allow free motion of the head without interference by the tip of the horn. This rigid process is present only in the female and although several entomologists had suggested that it received the ovipositor, its func- tion remained in doubt till Marchal ('06) showed that in the European Inostemma piricola, it really serves to receive the basal portion of the ovipositor which is much longer than the abdomen, so that when not extruded the base lies in the anterior part of the horn. The Inostemma studied by Marchal deposits its eggs in a Cecidomyiid larva which feeds within the small developing fruits of the pear. An examination of the North American Inostemma liorni Ashm. shows that, as might be expected, the mechanism of the oviposition is entirely similar to that of the European species. Marchal was at a loss to account for the apparent origin of the ovipositor within the basal tergite of the abdomen. Unfortunately the only specimens available have been mounted dry for a number of years, but dissections of these which I have made show that the ovipositor appears actually to arise within a n^embranous apical segment which has been invaginated so as to occupy the cavity of the process on the first segment. As there are six visible chitinized segments, this membranous tube is no doubt the seventh, or seventh and eighth abdominal segments and it must furnish the muscular apparatus for the manipulation of the ovipositor. The horn is therefore only secondarily a housing for the ovipositor. 1 The Australian genus Stylaclista Dodd ('15) is evidently very similar to Scorpioteleia, having the third to sixth abdominal segments produced into a long fleshy stylus. HYMENOPTEROUS PARASITES. 281 The genus Brachinostemma and several genera (e.g. Baryconus, Probary conns, Ceratoteleia, Caloteleia, Ceratobseus, etc.) of the closely allied family Scelionidre show a tubercle or very short horn arising from the dorsum of the first abdominal segment, but in no case does this ever attain a dcA-elopment approaching that seen in Ino- stemma. From the apparently rudimentary development of what is seemingly homologous to the horn in Inostemma, one might readily conclude that these genera show it in an incipient stage. From the standpoint of function this does not seem possible, however, as the projection is often so short that it does not serve to lengthen the space within the abdomen. Possibly the tubercle or horn may have been developed for some other reason and later served for the accommoda- tion of the ovipositor. The long horn appears to be unique, however, and no one has so far been able to attribute to it any other function. ^Ye may readily suppose that its ontogenesis is in direct response to pressure produced by the base of the developing ovipositor. It seems impossible that its length, at least in the incipient stage could be of any selective value, since most Hymenoptera provided with length- ened ovipositor have developed no structures or devices of any kind to permit of extensive retraction of this organ. After what has been said of the conditions prevailing in Dolichotrypes which have been considered at some length, it is evident that a more extensive knowl- edge of these minute Hymenoptera may lead to interesting conclusions concerning the relation between the morphology of the body and the function of oviposition. At the same time, it must be borne in mind that the horn of Ino- stemma does not vary to any excessive degree and that its form and size are at present as definite and clearly fixed in each species as are the other parts of the body, and that they are not variable like the abdominal segments of Dolichotrypes which have not yet attained fixed dimensions. 282 BRUES. Literature Cited. Ashmead, W. H. '93. A Monograph of the North American Proctotrypidte. Bull. U. S. Nat. Mus., No. 45, pp. 463, pis. 18. '97. Descriptions of Some New Genera and Species of Canadian Proctotrypoidea. Canadian Entom., vol. 29, pp. 53-56. Bateson, W. '92. On Some Cases of Variation in Secondary Sexual Characters, Statistically examined. Proc. Zool. Soc. London, 1892, p. 585. Brues, C. T. '09. A Preliminary List of the Proctotrypoid Hymenoptera of Washington. Bull. Wisconsin Nat. Hist. Soc, vol. 7, pp. 111-122. Brindley, H. H. '18. Notes on Certain Parasites, Food and Capture by Birds of the Common Earwig {Forficula auricular ia). Proc. Cambridge Philos. Soc, vol. 19, pp. 167-177. Brindley, H. H. and F. A. Potts. '10, The Effect of Parasitic Castration in Insects. Science, n.s. vol. 32, p. 836. Crawford, J. C, and J. C. Bradley. '11. A New Pelecinus-like Genus and Species of Platygastridse. Proc Ent. Soc. Washington, vol. 13, pp. 124-125, pi. 1. Dodd, A. P. '15. Australian Proctotrypoidea, no. 3, Trans. Roy Acad. So. Australia, vol. 39, pp. 384-405. '16. Australian Proctotrypoidea, no. 4, ibid., vol. 40, pp. 9-32. Felt, E. P. '18. Key to American Insect Galls. Bull. New York State Mus., no. 200, 310 pp. Giard, A. '94. Sur certains cas de dedoublement des courbes de Galton. CR. Soc. Biol., 1894 and Biologic Generale, pp. 335-338. Paris, 1911. KiefEer, J. J. '10. Family Belytidse. In Gen. Insect., fasc. 107, pp. 45. '16. Beitrag zur Kenntnis der Platygasterinse und ihrer Lebens- weise. Centralbl. f. Bakt., vol. 46, pp. 547-592. HYMENOPTEROUS PARASITES. 283 Marchal, P. '04. Recherches sur la biologic et developpment des Hymenop- teres parasites. Arch. Zool. Exper. et Gen., vol. pp. 257-335, 4 pis. '06. Recherches sur la biologic et le developpement des Hymenop- teres parasites. II, les Platygasters. Arch. Zool. Exper. Ser. 4, No. 6, pp. 485-640, pis. 8. Richardson, C, H. '14. Studies on the Habits and Development of a Hymenopterous Parasite, Spalangia muscidarum Richardson. Journ. Mor- phol. vol. 24, pp. 513-557, 4 pis. Winnertz, J. '53. Beitrag zu einer Monographic der Gallmucken. Linn. Entom., vol. 8, pp. 154-322. EXPLANATION OF THE PLATES. 286 BRUES. PLATE I. 1. Antennae of Polymecus (Dolichotrypes) minor sp. nov. 9 2. Antennae of Synopeas meridionalis sp. nov. 9 . 3. Antennae of Gastrotrypes spatulatus Gen. et sp. nov. 9 . 4. Antennae of Polygnotus simplex sp. nov. 9 . 5. Antennae of Platygaster tubulosa sp. nov. 9 . 6. Antennae of Tsdstasius crassus sp. nov. 9 . # Brues. — Hymenopterous Parasites. Plate I. ^^^^ ^„.,---^^ v/ Proc. Amer. Acad. Arts and Sciences. Vol. 57. 288 BRUES. PLATE II. 7. Wing of Polyrnecus (Dolichotrypes) minor sp. nov. 9 . 8. Wing of Synopeas meridionalis sp. nov. 9 . 9. Wing of Gastrotrypes spatulatus gen. et sp. nov. 9 . 10. Wing of Polygnotus simplex sp. nov. 9 . 11. Wing of Isostasius crassus sp. nov. 9 . 12. Wing of cf not associated definitely with any of the foregoing species. 13. Polyrnecus (Dolichotrypes) hopkinsi, 9 . (After Crawford and Bradley.) 14. Inostemma piricola Kieffer, 9 . (After Marchal). Brues. — Hymenopterous Parasites. Plate II. Ji^ N» ^ > =5 \ / i - Proc. Amer. Acad. Arts and Sciences. Vol. 57. VOLUME 56. 1. Kennelly, a. E., and Kurokawa, K. — Acoustic Impedance and its Measurement. pp. 1-42. February, 1921. $1.25. 2. Bell, Louis. — Ghosts and Oculars, pp. 43-58. February, 1921. $.85. 3. Bridgman, p. 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Brues, Charles T. — Some Hymenopterous Parasites of Ligoicolous Itonididfe. pp, 261- 288. 2 pb. May, 1922. $.85. 12. Thaxter, Roland. — A Revision of the Endogoneae. pp. 289-350. 4 pis. June, 1922. $1.25. ^ .* Proceedings of the American Academy of Arts and Sciences. Vol. 57. No. 12.— Jdne, 1922. CONTRIBUTION FROM THE CRYPTOGAMIC LABORATORIES OF HARVARD UNIVERSITY. LXXXIX. A REVISION OF THE ENDOGONEAE. By Roland Telaxter. With Four Plates. UV;^ A s^-^ CONTRIBUTION FROM THE CRYPTOGAMIC LABORATORIES OF HARVARD UNIVERSITY. LXXXIX. A REVISION OF THE ENDOGONEAE. By Rol.vnd Thaxter. Received March 17. 1922. Presented April 12, 1922. In preparing the present Contribution concerning the Endogoneae it has not been my intention to consider the subject in all its aspects, phylogenetic, cytological and other; and this revision has been under- taken chiefly with a view to the improvement of the systematic status of the family. For although relatively small, it has not escaped the taxonomic confusions and uncertainties which so frequently beset the path of the systematic mycologist, and it has seemed worth while to make at least an attempt to clear up some of the moot points relating to it, and at the same time to add such new information as I have been able to accumulate from personal observation or otherwise. I have therefore endeavored to obtain authentic information in regard to as many of the known forms as possible, and personally to examine as complete a representation of the type-material as could be assembled. Such value as this account possesses is therefore largely due to the courtesy of correspondents who have been so kind as to assist me in accomplishing these objects; and in this connection I desire to express my great obligation to Professor Abrams, of Leland Stanford, who has allowed me to examine all the Harkness types of Endogone in the University Herbarium: to the Abbe Bresadola, who has sent me a specimen of his E. reniformis collected by Rick in Brazil: to Dr. C. W. Dodge for Californian material collected by himself and by IVIr. H. E. Parks: to Professor E. C. Jeffrey for a very interesting collection from Little Metis, Quebec, given to me many years ago: to Professor G. Lindau for the privilege of examining portions of all the types of Hennings and Bresadola in the Berlin JMuseum; to IMr. C. G. Lloyd for a portion of his Endogone tuberculosa and other interesting forms; to Professor O. Mattirolo who has sent me for examination specimens of all his material of Endogone, including the types of E. Pamyaloniana and E. Tozziana; to M. N. Patouillard for confirming my determina- tion of his E. lignicola and for portions of the types of Acker mannia 292 THAXTER. Dussii and A. coccogcna; to Professor Carlos Spegazziiii for communi- cating the types of Endogone juegiana and E. argcutiua; and to jNIiss E. A. Wakefield for opportunity to see Berkeley's types of Endogone ansirolis and GlazicUa vesiculosa. Index. Ackermannia coccogena 333 Dussii 329, 333 Endogone arenacea 317 argentina 321, 324 australis 312, 313 borealis 318 canadensis . . . 317, 310, 318 fasciculata . . . 308, 309, 311 fulva 312, 317, 319, 320, 322, 326 fuegiana . . 303, 304, 309, 310 incrassata . 305, 293, 304, 316 lactiflua 306, 304, 308, 310, 312-314 lanata 306 lignicola 319 Ludwigii 298, 301 macrocarpa 312, 307, 315, 321, 326 malleola . . 323, 297, 322, 324 microcarpa 315, 297, 307, 323, 326 Moelleri 319, 320 multiplex 301, 304, 307, 317, 331 Pampaloniana . . . . 314, 313 pisiformis 298, 295-297, 304, 323, 327-8 pulvinata 319, 321 radiata 316, 305 reniformis 321 sphagnophila .... 299, 301 tenebriosa 314, 326 Torrendii 323, 324 Tozziana 326, 291 tuberculosa 293, 302, 303, 305, 311, 331 vesiculifera 309, 331 Endogone xylogena . . . 301, 300 Endogonella borneensis . . . 334 Glaziella abnormis ....... 338 aurantiaca 334 ceramichroa 338 splendens 338 sulphurea 338 vesiculosa 334 Glomus macrocarpus 312 microcarpus 315 Hypomyces alboluteus .... 334 Paurocotylis fulva 319 Protomyces xylogenus .... 300 Sclerocystis coccogena 333 coremioides . . . 328, 331, 332 Dussii 329, 328 pubescens 326 Sphaerocreas pubescens 326, 301 javanicum . . . 328, 327, 329 Dussii 329 coccogena 333 Stigmatella pubescens .... 326 Xenomyces ochraceus . . 328, 329 Xylaria aurantiaca 334 splendens 338 The fungi which are grouped in this assemblage of somewhat diverse forms are, in general, rather infrequently met with; owing in part to their apparent rarity, and partly to the fact that certain of the species, at least, are truly hypogaeous, and may develop at a depth of several inches below the ground, or beneath thick mats of Sp hagna or other mosses. While types of this sort are thus usually encountered by accident, or through the acquisition of what may be called a "hypogaeous instinct" which may enable one, after experience, to judge by various indications what situations are the most promising REVISION OF ENDOGONEAE. 293 for the collection of these and other fungi hypogaei, the recognition of others is surrounded by no such difficulty, since their fructifications may be developed free to the air, on mosses, rotten wood, leaves, dung or other substances above the leaf co\'er, or emerging from it. It seems not improbable that the vegetative hyphae of all the Endogoneae are at first continuous. In a majority of cases, however, the hyphae of sporulating conditions show at least occasional septa, which, in highly developed sporocarps like that of Glaziella, become very abundant. The spores developed from these hyphae are either zygospores, thick-walled acrogenous chlamydospores or thin walled spores formed endogenously in sporangia. The true relationships of the group to other families of fungi have long been a matter of conjecture, as is evident from the terms — asci, sporangia, cysts, vesicles etc. — which have been applied by various authors to the chlamydospores alone. But although the admirable researches of Bucholtz, who first (1912) published an account of its sexual reproduction, have thrown much needed light on its affinities, the group as a whole has been assumed to include forms of considerable diversity. The inclusion in a single genus of the zygosporic and chlamydo- sporic types, has hitherto been based entirely on a general resemblance in habit and habitat, and a similarity in the appearance of the two types of spore, and there has been no evidence which would indicate that the two were ever produced simultaneously in the same sporo- carp, or were closely associated in their natural habitats. This as- sumption proves, however, to have been justified; since in a single instance, among the northern forms collected by Professor Jeffrey and herewith described, zygospores and chlamydospores are so intimately associated in the same spore mass, that there can hardly be any question as to their specific identity. Although the zygosporic or chlamydosporic nature of these spores is usually manifest, except in very old material, it is not always easy in cases where they are surrounded by densely compacted hyphal tissue, to determine whether their origin is sexual or not, the elements involved being so compressed and distorted that conjugating processes, unless very conspicuously differentiated, might well escape notice. This is true for example in Endogone incrassata, Figures 17-19, or E. tuberculosa. There seems, however, at least in the genus Endogone, to be a rather fundamental difference between the two types. In the zygospores, which are usually surrounded by a more or less definite hyplial envelope, an outer wall is present, within which a continuous 294 THAXTEK. endospore is laid down, so that the contents is completely separated from the cavity of the origins. The contents is also more fatty and dense, often composed of distinct elements which may be very regular in size and shape (Fig. 9) and might even be mistaken for endospores. The chlamydospores, on the other hand, although' they may closely resemble the zygospores, do not appear, as far as I have seen, to pro- duce a continuous endospore; unless the otherwise anomalous Glaziella in which such an endospore is clearly distinguished (Fig. 91) proves to be an exception. For this reason, in a majority of species, the proto- plasm of the chlamydospore and that of the sporophore are continuous, being connected by a protoplasmic isthmus which may remain un- broken even in mature spores (Fig. 46), or may be finally pinched off in the middle by the gradual thickening of the lateral walls. In a smaller number of instances, the separation between the cavities of the spore and sporophore is accomplished at an early stage through the formation of an independent septum, Figures 52-59 and Figure 85. Spores of the latter type have, for the most part, much thinner walls than those of the former, and were regarded by Bucholtz as perhaps young sporangia. An examination of various species and copious material, however, has convinced me that they are homologous with chlamydospores of the first mentioned type. The so-called sporangia which have been above alluded to. Figures 60-78, which have been associated only with the genus Endogone, are quite unlike the other types of reproduction; and although I have followed previous writers by including them in this genus, there is no evidence beyond a certain resemblance between the sporangiocarp in the one case and the sporocarp in the other, which would tend to confirm the correctness of this reference. These sporangia are termi- nal vesicles, formed in a solid mass at the extremities of branching sparingly septate filaments which radiate more or less definitely from a cushion-like base. The spores which they contain are variable in size, form and number, thin-walled, with dense or fatty contents, and result from a total cleavage of the sporangial protoplasm. They are so characteristic that it would be quite impossible to mistake them for the spore-like masses above mentioned which may occur in zygospores or sometimes even in chlamydospores. Baccarini (1903) was of the opinion that these sporangial types should be removed from the Endogoneae and placed in the Mortierel- leae; the sporangium in both cases being separated from the sporangio- phore by a simple septum. As Bucholtz remarks, this disposition appears to be somewhat premature. It must be confessed, however. REVISION OF ENDOGONEAE. 295 that if these sporangial forms are rightly inchided in the Endogoneae, it seems very probable that the two families should be regarded as very closely related, at least; since they are similar in two other important characters; namely, through the production of specialized zygosporic envelopes, and the presence of highly specialized acrogenous chlamydo- spores. As far as I am aware, there has as yet been no successful attempt to germinate the spores of any of these fungi, or to grow them under artificial conditions; and in my own experience I have been unable, after repeated attempts, to induce the zygospores of Endogone pisi- formis Lk. {sphagnophila Atk.) to germinate; or to procure any characteristic growth when uncontaminated spore-masses have been transferred to agar nutrients. The spore-masses of this species, when wintered over out of doors, have also failed to develop further. When placed on fresh sphagnum in a moist chamber for a protracted period during the summer, they usually l>ecome covered by a thin white coating of nondescript hyphae: but although various peculiar Zygo- mycetes, to which reference has been made in a former paper, (Thaxter (1897) p. 12) have at times been observed in such cultures, there is no reason to believe, even though, as in some instances, they seemed to grow from the masses themselves, that the association was other than an accidental one. It seems very probable that the thick-walled spores of the Endogoneae, as in various other instances, germinate as a rule only after special preparation, or under special conditions, and that in Nature they are eaten by various animals; continuing their development after being voided. This is suggested by the fact that I have myself observed uninjured spores of species of Endogone in the stomach-contents of shrews and of myriopods. Until successful cultures have been made, and the development of the three spore types has been successfully followed, or at least until more careful and extended field observations have given some evidence of their actual connection it cannot be assumed that they should all three be included within the limits of a single genus. The literature of the Endogoneae, since the type species of the genus Endogone was described by Link in 1809, has been scattered and not very voluminous. With the exception of the paper by Buc- holtz, above mentioned, and the enumerations in the Kryptogamen- flora of Rabenhorst and of Cohn and the Pflanzenfamilien, there has, I think, been no general summary even of this genus. Von Holinel in his Fragmenta, Nos. VI, X and XV, discusses the synonymy and relationships of the genera Endogone, Endogonella, Sclerocystis, 296 THAXTER. Xenomyces, Ackermannia and Sphaerocreas ; and numerous other references to the genus Endogone, or descriptions of new species, are to be found here and there in various other pubhcations. In the appended list of literature, however, only such titles are included as are in some measure essential, and those which have reference merely to records of occurrence have been omitted. Turning first to the genus Endogone, and following the conception of the genus which has been adopted by Bucholtz and all recent writers, one is forced to include in it all the three categories of spore- forms above enumerated, namely, zygospores, sporangia and chlamy- dospores. In order to avoid new names and combinations I have adopted this procedure as a provisional solution. It may be well to repeat, however, that although the sporangial forms arise in general from a similar vegetative body, and are associated in somewhat similar aggregations in similar habitats, their connection with the other types has not been definitely indicated, even by close association in nature, and their inclusion in the same genus is based on a pure assumption. Whether it may prove desirable to retain the name Endogone for the sexual and chlamydosporic forms and to apply a different name to those which form sporangia is not as yet clear. It may further be pointed out that the presence of isogamy and of heterogamy, of specialized spore envelopes or their absence, as well as of simple and multiple aggregations of the zygospore masses, may similarly lead to a subdi^■ision of the sexual forms themselves, under more than one designation. The desirability or the reverse of either of these pro- cedures will, however, doubtless become more clear as the lacunae in our knowledge of the group are gradually filled. The reasons which have determined the selection of the sexual forms as the true representatives of the genus, as originally founded, are based on an examination of the original figures and description given by Link (1808), of the type-species, Endogone pisiformis, on p. 33 of the apparently rare publication in which his paper is contained. The exact wording of this description is as follows : "38. Endogone. Sporangium subglobosum, extus floccosum, intus grumosum sporangiola minuta,globosa, membranacea,sporidiis repleta. "Praecedenti generi affine, (Tuber), supra terram in muscis crescit hypothallo radiciformi. ISIembrana externa sporangii tenuis floccosa. Contextus caeterum vesiculosus, microscopio simplici inspectus grumosus, at compositi ope conspiciuntur sporangiola, ut in prae- cedenti genere, dispersas inter vesiculos multo minores. Sporidia minuta, globosa, sporangiolis inclusa. Unica species. J REVISION OF ENDOGOXEAE. 297 "E. pisiformis, irregulariter glohosum, lutescens membrana floccosa inductum. IMagnitudine pisi. Fil)rillis panels muscis adnascitur in silvis abietinis. jMenil)rana floccosa inducta tenuissiina, sporangium intus colore lutescente Tuberis, at non venosum, sed grumoso granu- losum. Segmenti transversalis particulam, V. fig. 52a, sporangiola cum sporidiis ibid. lit. b." The figure " a " referred to, shows a portion of the spore-mass covered by a radiating sterile tomentum (membrana floccosa tenuissima) of tapering filaments, evidently more or less diagrammatically repre- sented. The spores, which are shown embedded in the general mass (sporangium), are not subspherical, but more nearly elliptical, with the exception of those which may be assumed to be viewed end on. Figure " b" shows several of these spores (sporangiola) which have been forcibly and irregularly broken, as is evident from the rent through which the contents is represented as emerging. This contents is made up of granules indicated by single black dots, the "sporidia rninuta" of the description, which bear no resemblance to resting spores and could not by any stretch of the imagination be regarded as intended to represent the large thin-walled spores of the sporangial type. This description is sufficiently clear, although, like most descriptions, in- complete, and taken in connection with the figures, which are not bad for the period, afford a reasonably satisfactory basis for determination. Since K. jyisiformis is the generic type, it is a matter of much im- portance to determine with some approach to accuracy, to which of the European forms now recognized it may be assumed to corre- spond. Bucholtz, who may have seen transcriptions, only, of the original paper, and may have been misled by the confusing use of the terms sporangia sporangiola and sporidia, has assumed that- the classic specimen collected near Naples by Vittadini and distributed in the Fungi Europaei No. 2516 under the name Endogonc microcarpa, was to be regarded as the true pisiformis. It seems cjuite impossible, however, to reconcile the characters of the Mttadini form, which is the Endogonc mallcola of Harkness, with the account given by Link whose figures alone are sufficient to preclude the possibility of such a con- clusion. The more important points brought out by Link's account indicate that he was dealing with the type of sporocarp usually found in Endo- gonc, consisting of yellow ellipsoid thick walled spores with coarsely granular contents, associated with smaller vesicular structures, and irregularly disposed in a solid compact rounded mass surrounded by 298 THAXTER. a rather conspicuous "thin floccose membrane," and developed above ground on mosses. If one compares with this account the characters of the other known European types, none seem to correspond so closely as E. Ludwigii Bucholtz (E. sphagnojMla Atk.) No other species is found, as far as I am aware, growing on mosses above the surface of the ground, while its yellow ellipsoid spores with uniform coarse granular contents, and its conspicuous thin white superficial tomentum further distinguish it. The vesicular swellings of its hyphae, which are sometimes conspicuous among the larger spores, may further correspond to the "vesiculae multo minores" of Link. Since for the reasons above indicated the reference by Bucholtz of E. maUeola Hark, to E. pisiformis Link cannot be regarded as a possi- ble solution of the difficulty, and since it is quite necessary to form some reasonably plausible opinion as to what constitutes the Type of the genus, I have felt it desirable to follow Krieger (1902) and the earlier opinion of Bucholtz, in referring to E. pisiformis Link the spe- cies more recently named by the latter (1912) Endogone Ludwigii. ENDOGONE Lk. Link (1809), p. 33. Ghmus Tulasne (1845), p. 63. Hypogaeous or epigaeous: producing thick-walled isogamous or heterogamous zygospores with or without specialized envelopes: thick walled acrogenous non sexual chlamydospores: or thin-walled sporangia. The three types, as a rule, produced separately in com- pact groups, which may be single or associated in a common mass, naked or surrounded by a variably developed pseudoperidium or tomentum, and may form either a definite sporocarp or an indefinite loosely coherent spore-mass. Type Species. Endogone pisiformis Link. (Figs. 1-7.) Link (1809), p. 33, Taf. II, fig. 52, a & b. Bucholtz (1902), p. 81, Tab. II, fig. 13 and V, fig. 4. Krieger (1902), Fungi Saxonici, No. 1651. REVISION OF ENDOGONEAE. 299 Endogone Lmlwigu Bucholtz (1911), p. 194, Taf. IX, figs. 77-87. E. sphagnophila Atkinson (1918), p. 16. E. xylogena Schroeter (1887), p. 260, nee. Saccardo (1877), p. 1-4, sub Proto- myces. Thaxter (1897), p. 12. Spore-masses waxy when fresh, horny when dry, pale to golden yellow, becoming somewhat orange yellow, subspherical to reniform, or lobed, less often convolute, flattened, umbilicate below: covered by a thin tomentum, clear white when dry, formed by characteristic, thick-walled hyphae 4-6- m in diameter with numerous free, projecting, distally attenuated branches. The substance of the spore mass con- sisting of an irregular plexus of stout branching non-septate filaments, showing numerous irregular vesicular enlargements, becoming more or less obliterated as the irregularly crowded, broadly ellipsoid to ovoid, thick-walled, pale orange yellow zygospores mature. Spore- masses (dry) 2-7 X 1-2 mm. thick. Zygospores, 35-60 X 30-45 fx, the wall subhyaline 3-5.5 jj. thick. Peridial hyphae X 3-8 fx. Usually above, rarely below the leaf cover; on mosses, especially near the tip of Sphagnum; on leaves, twigs, dung, rotten logS; etc., in moist situations, especially in coniferous woods. Tempei'ate Europe and North America. This species is without doubt very generally distributed in temper- ate America; since it is already known to occur in Maine, New Hamp- shire, Connecticut and eastern Tennessee (Thaxter); West Virginia (Sturgis); New York and Maryland (Atkinson), and in Michigan (Kauffman). In my own experience it has proved not at all uncom- mon, and was first met with at Kittery Point, Maine, in 1886, when young conditions, showing the early stages of conjugation were ob- tained. Although it is found most frequently at or near the tips of Sphagnum, especially in moist coniferous woods, and is conspicuous in this position from its bright color, it bears no definite relation to this substratum as a host; since it occurs also, as above indicated, on various other substances. Its waxy consistency, when fresh is, as noted by Schroeter, characteristic; as is the hard almost horny char- acter of the dry spore mass, which loses its bright color, becoming dirty yellowish; the variably developed superficial tomentum assum- ing a more noticeable clear white appearance, owing probably to the refractive character of the thick walled filaments which compose it. The size and form of the spore-mass varies considerably from nearly round to flattened and somewhat convolute. The largest individual seen measures 7 mm. in width when dry. The early conditions of development are much more difficult to 300 THAXTER. detect, from their small size and much paler color. The process of conjugation is not progressive in the developing mass; but occurs almost simultaneously throughout it, the rather rapid enlargement of the whole being due to the simultaneous increase in size of the indi- vidual zygospores. The gametes are subequal, and do not differ from one another more than is frequently the case in other isogamous types. They are subcylindrical and lie parallel to one another, distinguished by a clean cut septum at some distance below their adherent tips. Figure 1. The developing zygospore rises from this point of contact, above and between the extremities of the gametes, Figures 2-6. The successive stages in this process are not unlike those figured by \ an Tieghem (1873), PI. Ill, figs. 88-93, in Synccphalis cornu. Before full maturity, the hyphal elements of the mass are con- spicuous, and rather characteristic from their large size, their branching and the development of vesicular swellings which I have assumed to be the "vesiculi multo minores" mentioned by Link, and which are referred to by Bucholtz as " stellenweise verbreiterungen." As the zygospores mature, these elements become compressed between them, and may be hardly recognizable, their flattened remnants forming, in many cases, an irregular envelope about the individual spores. The branching terminations of the filaments which form the super- ficial tomentum are well figured by Bucholtz (1911), fig. 77, and possess great individuality, Figure 7, but are not always conspicuous in older individuals. The prominence of this tomentum varies greatly in different individual masses, and under different conditions. It seldom seems to be so copiously developed as is represented in the figure of Link, which is evidently somewhat diagrammatic, and in older specimens may appear to form a rather even covering of appar- ently nearly uniform elements. The description given by Schroeter of Endogone xylogcna corre- sponds so closely to this species, that I have included it as a synonym. It seems quite improbable that the plant which he examined could have been the Protomyccs xyhgenns of Saccardo; since the latter is without hyphae, and corresponds in all respects to the sclerotium- condition, "Phylloedia," of some myxomycete: its habitat, buried in soft rotten wood and exposed only by the weathering of the latter; its yellow color, and the general appearance of its spores, being the same. The figures given by Saccardo (1877) in the Fungi Italici, fig. 104, show the somewhat irregular outline and the characteris- tically thickened, but ill defined, walls of this well-known condition of the myxomycete plasmodium. REVISION OF ENDOGONEAE. 301 With reference to the occurrence of this species in Europe, it may be mentioned that the single specimen collected by Bucholtz in Livonia was found " in einem nadelwald unterirdisch," and was associated with insect-remains, which suggests that it may have grown on the dung of some small animal, a habitat which I have myself observed. The apparently copious material collected in Thuringia by Ludwig, which forms the basis of the account given by Bucholtz, was found on the dung of Liparis caterpillars. The specimens distributed by Krieger were found "Auf Moos, faulenden Bliittern, Aestchen, unter Strau- chern von Vaccinium myrtilus auf dem Fichtelberge in Erzgebirge." With regard to mutual identities in connection with this species, it should perhaps be clearly stated that while the use of the name E, pisiformis and the inclusion of E. xylogena as a synonym represent merely my personal conclusions. Professors Atkinson and Bucholtz have both examined the material on which the present account is based, and have pronounced it identical with E. sphacjnopkUa in the one case, and E. Liidwigii in the other. It may further be mentioned that one of the specimens distributed by Krieger, has been examined by me personally, and is also identical; although a second specimen in the same copy of this set, the gross appearance of which is very similar, proves to be Sphacrocreas pubesccns. As it is stated that the fungus was found " sehr selten," it may be assumed that the distribution is a miscellaneous one, accumulated from more than one gathering. The possible relation between Sphacrocreas pubesccns and Endogone pisi- formis will be further alluded to under the former species. For convenient comparison, the description of E. xylogena given by Schroeter (1. c.) may be here appended. "Endogone xylogena (Saccardo (1877): Protomyces x.). Fruiting bodies irregularly rounded, flattened, 3-4 mm. broad, 1-2 mm. thick, waxy when fresh, horny when dry, reddish yellow. Peridium thin, formed from 3-5 yu thick, strongly refractive hyphae, smooth. Gleba homogeneous, consisting of closely woven hyphae between which the spores are disposed. Spores spherical to elliptical or ovoid, 35-50 X 26-40 fx, the wall 6 fi thick, nearly hyaline, contents clear orange yellow. Endogone multiplex nov. sp. (Figs. 8-10.) Fruiting body about 15 X 12 mm., dirty whitish, turning yellowish brown in alcohol; somewhat lobed, the surface rough from the pro- jecting contours of the very numerous small, more or less firmly 302 THAXTER. coherent, rounded or somewhat irregular spore-aggregates, of which the mass as a whole is composed, and throughout which a large amount of finely divided humus material is incorporated. Individual spore- groups more or less rounded, or somewhat irregular, mutually coherent, or readily separable, 350-700 jj. in diameter, and including from ten to fifty spores each, more or less; each group surrounded by an envelope of hyphae among which a considerable amount of humus material is incorporated; the hyphae variable in diameter, 4-18 /x, thick-walled, rather brittle, freely branched, three or sometimes four branches often radiating from subtriangular or angular enlargements, especially in the larger ones, which are rather conspicuously distinguished, though scanty. Zygospores yellow, spherical, oblong to ovoid or piriform, often irregularly subangular from pressure, 80-90 X 60-84 ^t; the endospore clearly defined, slightly yellowish, about 5 yu; the exospore hyaline and, when freed, swelling to 8-10 n; the contents rather bright yellow, composed of nearly spherical fatty bodies 4-8 /x in diameter which completely fill the cavity. The attachments of the suspensors clearly defined, sometimes approximated, more often distant: the spore surrounded by a clearly defined, relatively thick, separable envelope, 8-12 p. thick, of closely felted hyphae. Growing beneath the leaf cover beside a path in mixed deciduous woods (oak and hickory) on Cutts Island, Kittery Point, Maine: September 15, 1902. This species is most nearly related to E. tuberculosa, but differs in various essential points. The individual spore-masses are, as a rule, very readily separable, so that a small fragment of the fruiting body, when teased or rubbed under the cover glass, separates to a mass of rather uniform coarse granules, which represent the individual spore- groups. Figure 10: the envelopes of which are composed largely of humus particles which often wholly conceal the spores within. The material is unfortunately fully matured, and it is thus impossible to determine the exact nature of the process of conjugation, and even the suspensors are for the most part disorganized to such an extent that their form and limits can no longer be made out. The relation and attachment of the latter to the spore are very characteristic. They are always quite distinct. Figures 8-9, sometimes close together, but usually separated by a considerable interval; in this respect re- calling the similar relation so often seen in the zygospores of Choane- phora. On treatment with potash, the separable exospore and the surrounding filaments become considerably swollen and gelatinous, so that their limits are determined with difficulty. REVISION OF ENDOGONEAE. 303 The peculiar characters of this species illustrate the culmination of the tendency toward a definite grouping of the spores within the gleba, which is present to a less marked degree in E. tuberculosa and E. fuegiana. The sexual nature of the spore-origin is unquestionable from the two distinct origins are present in all spores. The alterna- tive that they may be intercalary and represent a lateral bulging, so to speak, in the continuity of the hypha, is an explanation which is rendered quite improbable by our knowledge of spore-formation in all the chlamydosporic types. The conjugation is evidently somewhat peculiar, as is evidenced by the often remote origins, and it is to be regretted that, owing to the fact that the whole spore-mass is hardly distinguishable from a slightly coherent mass of earth, the younger stages are not likely to be found, unless by accident. Endogone tuberculosa Lloyd. (Figs. 11-16.) Lloyd (1918), p. 799; fig. 1239. This species has been described and its gross appearance well illus- trated by Lloyd, to whom the writer is indebted for a small portion of the type material on which the following notes are based. It was col- lected in New South Wales by Mr J. B. Cleland, who states that it was found just at the surface of the ground, apparently partly buried in it, if one may judge by the coating of earth which completely envelopes it. Its gross characters are peculiar from the fact that the gleba is not a continuous and undifferentiated spore-mass, but is in a sense compound. The sporogenous area, which is only visible in sections. Figure 11, is very irregular in outline, pushing indeterminate lobes or extensions outward into the surrounding covering of earth, which thus varies greatly in thickness, and appears to be held together by a scanty penetrating mycelium. It is possible, after slightly moistening the cut surface, to determine that the golden yellow spores are arranged in rounded masses of variable size and shape, or are associated in larger somewhat less definite areas. In either case they are often, though not always, separated by intruding layers of the earthy matrix, the presence of which is indicated by its darker color, and which may be even more intimately incorporated in the general mass, although none appears to occur within the individual spore-groups. In these spore-groups, or areas, the more clearly defined of which 304 THAXTER. may be from 350-1000 fi in diameter, more or less, the bright yellow spores are closely packed and coherent, each surrounded by a thin, but as a rule clearly defined, envelope of closely matted finer hyphae. Penetrating the larger groups or areas, or separating the smaller ones, vein-like wefts of coarser filaments, forming an irregular pseudo- parenchyma, may be present. Figure 12, so that the general appear- ance of the cut surface is not unlike that of one of the Tuberaceae. The individual spores. Figures 12-16, are often irregular from pres- sure, and very variable in size and outline; subspherical or more often longer than broad, elliptical, subpiriform or often elongate, 50 X 42- 150 X 90 fjL, the average about 90 X 65 ^t; the exospores about 5-6 fx, becoming very thick, even 15 ju; t^e endospore comparatively thin, about 1-2 fjL. The yellow contents consists of not always dense, granular fatty protoplasm, usually associated with larger fatty masses or globules; but in certain fully mature individuals, it appears to have lost its color, becoming hyaline; while the exospore is greatly thick- ened, Figure 14, intruding irregularly, somewhat as in E. incrassata, and throwing the endospore into irregular folds. Although, owing to the mature condition of the specimen, the spore- origins are for the most part shriveled or destroyed when freed from the tenaciously adherent spore-envelopes, a sufficient number have been isolated to satisfy me that two hyphal elements are involved in spore-production, which are associated and differentiated much as in E. lactiflua; although relatively smaller and less conspicuously difPer- ent, one from the other, than in this species. In one instance, only, Figure 13, has it been possible to determine with some exactness the more normal appearance and relation of the two conjugating elements, although many have been observed in which the remains of corre- sponding structures were clearly traceable. In the type figured by Lloyd, the surface of the specimen is con- siderably and irregularly roughened, pitted or lobed, the roughness having apparently suggested the specific name. This tuberculate habit does not, however, appear to be related to the presence of the characteristic spore-groups, and is merely a modification of the earthy covering. The species is more like E. pisiformis in the form and color of its spores, but resembles E. lactiflua in its type of conjugation. In the grouping of its spores and its yellow color it recalls E. viuUiplex, which is nevertheless readily distinguished by the two discrete suspensor- insertions which characterize this species. The grouping of the spores is similar to that found in E. fuegiana, which, however, forms REVISION OF ENDOGONEAE. 305 a compact continuous spore-mass, without incorporated foreign material, and in which the origin of the spores and spore-groups is quite different and apparently non-sexual. Endogone incrassata nov. sp. (Figs. 17-19.) Fruiting body even or somewhat lobed, yellowish, with a whitish scaly or reticulate crust variably developed, about 2-5 mm. in diame- ter when dry. Gleba firm and compact, yellowish; the hyphae thin- walled and vesicular, or running in strands or bundles between the spores; the thin peridial region of more slender thick-walled filaments. Spores scattered thickly, without definite arrangement, throughout the mass of the gleba, which contains no foreign matter; more nearly isodiametric, somewhat irregular in outline, subspherical to broadly oblong, at first filled with rather uniform yellow subspherical fatty granules, about 3-5 /j., the continuous endospore clearly defined, thinner than the exospore; the two about 8 /x thick; the exospore becoming much thickened, 16-20 /jl, intruded toward the center and pushing the endospore into folds, the contents losing its color and granular character. The spores 66 X 64-75 X 85 /x. Under spruce, about two inches below the surface of the cover; with a distinct aUiaceous odor. Gerrish Island, Kittery Point, Maine; August, 1896. Three specimens of this species were found associated, and close by a single individual of E. radiata, of which it may possibly prove the sexual form. The gleba is so dense, and its elements surrounding the spores so vesicular, that it has been impossible to make out with cer- tainty the character of the gametes which are evidently small, not clearly distinguished and almost obliterated by the enlargement of the spores and the consequent pressure. In a few instances, appearances have been seen such as are represented in Figures 18-19; but, in the dense pseudotissue about the spore, it is quite possible that the appar- ent conjugating spore-origin may be in reality due to an accidental juxtaposition of gleba elements, bearing a superficial resemblance to conjugating structures. The spores when fully matured. Figure 19, resemble those of E. tuberculosa, Figure 14, although the wall of the exospore becomes rela- tively thicker and the endospore is thrown into deeper and more complicated folds by its intrusion. In this condition it is quite hya- 306 THAXTER. line and impenetrable by stains, the contents losing its granular char- acter entirely. The spore-envelope is thin and not clearly differ- entiated. The scaly or flecked appearance of the surface of the sporocarp is due to patches of loose hyphae which project from the peridium, and in section appear as flat tufts. Endogone lactiflua Berkeley (1846). (Fig. 20.) Berkeley (1846), p. 81. Tulasne (1862), p. 183. Bucholtz (1912), p. 155, figs. 1-61. Endogone lanata Harkness (1899), p. 280. This species has become for the first time thoroughly well known through the researches of Bucholtz, who was not only the first to see and to describe the sexual origin of its spores, but to figure clearly the remarkable envelope which surrounds them at maturity, formed from labyrinthine filaments which eventually become thickened and modi- fied to form what he has called a "flammenkrone," which is firmly adherent to the exospore. Both the envelope and the flammenkrone, however, vary, as is mentioned by Bucholtz, (1912), p. 165, in different individuals, apparently according to the age of the spore-mass, and in some of the Hesse specimens in the Farlow Herbarium neither are striking or easily recognized; while in others they are apparent at a glance. The same is true of material which the wTiter has collected at various times and in various localities in New England; at South Billerica, Mass.; at Kittery Point, Maine, where seven different gatherings were made; and at Intervale, New Hampshire. In all these gatherings, which were mostly of single specimens, the gross size is smaller and the spores themselves larger than in the Hesse speci- mens; and while in some the labyrinthine envelope-filaments (Buc- holtz, fig. 50), though finer, are quite as distinct and the fiammen- krone clearly distinguished, in a majority of cases these structures are not clearly visible, except that a well developed hyphal sheath is always present. Entirely similar conditions are, however, seen in some of the Hesse specimens, so that it seems probable that their distinctness may be a matter of age or some of the circumstances associated with their growth. Although in the Hesse material the spores are usually only 100 /x in diameter, while in the American they REVISION OF ENDOGONEAE. 307 are 120-125 fi, specimens received from Hesse by Ed. Fischer are reported to be 115-125 X 70-90 i^l, and in the large number of cases reported by Bucholtz, the range of variation is 68-160 X 60-104 fj.. The discrepancy is thus not so great as it might at first appear; al though further examination may indicate that more than one specific form is represented in this series. Although the occurrence of this species in America has not been hitherto recorded, it appears to have been collected several times by Harkness in California. Through the courtesy of Professor Abrams of the Leland Stanford Herbarium, I have had an opportunity to examine all the material of Endogone referred to by Harkness, (1899), in his paper on Californian Hypogaeous Fungi, including " E. lanata" sp. nov., " E. microcarpa" Tul. and " E. viacrocarpa" Tul. The portions of these specimens communicated are similar in color and appearance, and it would be impossible to distinguish either of them by their microscopic characters from the eastern material above re- ferred to. In all, the conjugating processes are clearly defined, and the spore-envelope well developed. In the specimen marked " E. macrocarpa" this is especially true, the flammenkrone, though not as striking as in the best developed Hesse specimens, being clearly present. The size of the spores in these Californian specimens is also similar, the longer axis varying from 125 ^t or less to 160 /z: a range similar to that reported for the European types. In a single specimen found at Kittery under beech trees, the gleba is dark blackish brown, the color being apparently due to the fact that a large amount of finely divided humus material is incorporated throughout its substance, a condition seen elsewhere in E. multiplex and a few other species. The zygospores differ somewhat in possess- ing a somewhat roughened, smoky brown exospore, distinctly unlike the yellowish wall of the ordinary type. It has not seemed desirable to separate this form specifically, however, on the basis of a single specimen. For further details in regard to E. lactiflua, the admirable and very complete account of Bucholtz should be consulted. The possibility should be borne in mind that the very variable series of forms now included under this name may prove to represent more then one species, when they become more thoroughly known, and their life- histories have been traced. In the present state of our knowledge, however, the use of a single nanje to designate them seems in every way desirable. 308 THAXTER. Endogone fasciculata nov. sp. (Figs. 21-28.) Spore-masses spongy, loosely coherent, rather thin and irregularly lobed, somewhat amorphous, 10-14 X 4-5 mm., but very variable, incorporating more or less of the substratum (Sphagnum) and other foreign matter. Chlamydospores in rounded or somewhat elongate or irregular coherent groups, associated with less definitely distin- guished masses of readily separable zygospores; pale yellowish or faintly brownish, mostly spherical or somewhat longer than broad, 60 X 60-85 X 70 fx, the wall becoming relatively very thick, 6-10 fx. Zygospores immature, irregularly spherical, colorless, about 50 n, arising from the larger of two unequal gametes. In Sphagnum. Little Metis, P.Q. E. C. Jeffrey. This species is in some respects the most interesting member of the genus, since it is not only peculiar from the grouping of its spores, but presents the only instance in which zygospores and chlamydospores have been found intimately associated in the same spore-mass. It thus furnishes the first indubitable eA'idence that the zygosporic and chlamydosporic types have been rightly included in a single genus. None of the zygospores examined are mature, but there is no indi- cation that any special envelope is developed about them, as in E. ladiflua and some other sexual forms; although the process of forma- tion. Figures 23-26, is very similar to that which occurs in the last mentioned species. The hyphae with which they are associated are thin-walled, scanty and evanescent; so that even in the youngest stages of conjugation, the exact origin and relation of the progametes is not clearly evident. Although this cannot be regarded as deter- mined beyond question, examination of young stages under an immer- sion seems to show that the type of conjugation is homothallic, and that the progametes arise in proximity to one another from the same filament. The gametes are distinguished much as in E. ladiflua, one being larger than the other, and bearing the zygospore, which bulges upward; both remaining attached, with slightly thickened walls and septa. The groups of zygospores are more irregular and undifferen- tiated than those of the chlamydospores, among which they are irregularly distributed in continuous masses. The chlamydospores arise from a plexus of clearly defined, thick- walled, variously bent and interlaced branching hyphae, which form a core from which short irregular sporiferous branches grow radially REVISION OF ENDOGONEAE. 309 outward. The chlamydospores are thus at first rather firmly asso- ciated in grape-like clusters, which may be of definite rounded outline, Figure 21, or longer or more irregular. This definite relation seems to be more or less obscured in older specimens in which the hyphae tend to break up, as in other species of the genus. It should be mentioned that zygospores do not seem to be invariably associated with the chlamydosporic form. The chlamydospores themseh'es are rather uniform, commonly more or less spherical or but slightly longer than broad, and when fully mature possess a relatively very thick wall, surrounding a coarsely fatty contents. The species is most nearly related to E. vcsicuUfcra, which seems very clearly distinguished by the peculiar clavate empty vesicles which are associated with the chlamydospores. In the grouping of its spores it also bears some resemblance to E. fuegiana, which is at once distinguished by its hard continuous gleba. Endogone vesiculifera nov. sp. (Figs. 29-32.) Spore-mass loose in texture and without definite form, about 5-8 X 4 mm., incorporating more or less of the substratum (Sphagnum) and some other foreign matter. Chlamydospores arising in groups, rounded or more elongate, often nor clearly defined; pale yellowish, spherical or slightly longer than broad, rather uniform, about 65 X 05 (JL, the larger 80 X 70 yu: arising from fascicles of intricately woven, branching, thick walled hyphae, and borne terminally on short radiat- ing branches; associated with broadly clavate vesicular cells, 100-125 X 50-64 jj., which extend outward beyond them. In Sphagnum, Little Metis, P.Q.' E. C. Jeffrey. The material of this form is somewhat scanty, although sections of three different individuals are preserved. It resembles E. fascicvlata very closel}', the chlamydospores being very similar in size and shape and similarly grouped about a core of thick-walled hyphae. It is readily distinguished, however, by the presence of numerous pear- shaped or broadly clavate, nearly empty, thin-walled, sterile vesicular structures which arise in company with the chlamydospores from slender short branchlets. These bodies are very characteristic, and although their origin is the same, are by no means ordinary chlamydo- spores which have failed to develop. They arc no doubt the homo- logues of spores, but cannot be directly compared with the numerous 310 THAXTER. empty abortive vesicle-like spores which are conspicuous, for example in E. canadensis. In many cases their broad projecting terminations form a continuous margin about the spore-groups. Those of the latter which are peripheral, may be further enveloped externally by a closely woven layer of fine, thin-walled, hyphae, which may penetrate inward to some extent, between the vesicles and spore-groups, entering the spores themselves and filling them more or less completely. This parasite seems similar to that which attacks E. lactifliia, E. fuegiana and other species. Endogone fuegiana Spegazzini. (Figs. 33-34.) Spegazzini (1887a), p. 6, No. 5; (1887b), p. 120. Through the courtesy of Professor Spegazzini I have had an op- portunity to examine the type of this species collected on Staten Island, Straits of Magellan. In its present condition the type does not show all the characters mentioned in the original description which, since the publication in which it appeared is rare, should perhaps be quoted in extenso. "Globoso vel elliptico repanda, extus alba, levis vel vix sub lente valida flocculosa, parvula (2-5 mm. diam.), inferne saepius umbilicata vel depresso-rugulosa centroque nodulosa vel subcicatricosa, uda compactiuscula tenacella; sicca dura, fere cornea: cutis carne arete adnata persistens; caro sordide alba sub sectione fulvo-maculata, ob punctulos rufos dense congestos: puncti 7-8 cellulares, globoso sub- polygoni (180 ^i diam.), carne innati, nunquam confluentes: cellulae punctulorum sphaeroideae e mutuo pressione saepius ovoideae (80 X 65 fx) laeves, crasse tunicatae ad verticem precipue, inferne subapicu- latae ac nodulo majusculo obscuriore donatae, fulvae vel subtestaceae. Inodora, insipida." Found under moss on Staten and Clarence Islands, Straits of Magellan. There has been some question as to the true position of this species, owing to the characteristic arrangement of its spores, the "cellulae" of the above description, which are more or less definitely and com- pactly associated in small groups of six or usually more, Figure 33, separated by variably distinct strands of compact parallel hyphae, an arrangement which gives an irregular and rather faintly areolate appearance to sections of the gleba. This has led to the suggestion that the plant might be an immature condition of some tuberaceous » REVISION OF ENDOGONEAE. 311 form. The species Is, however, a well defined Endogone. The spore- groups -are smaller and more clearly defined than those of E. tubercu- losa. Spegazzini remarks that the spore-groups are never confluent; but a section from the dried material shows that they are not always distinguished with great clearness, and are at least often in close contact. The spores, unlike those of E. tuberculosa, are reddish brown, con- siderably smaller and more nearly spherical, though usually irregular from mutual pressure. Their greatest diameter seldom exceeds SO /x, while that of E. tuberculosa is often as much as 125 m- The gleba is a dirty brownish yellow with a reddish tinge, horny when dry, the strand which separates the spore-groups, which are not always clearly marked, having a darker brownish color. The gleba, unlike that of E. tuberculosa, is continuous in the sense that, as far as I have seen, it contains no incorporated foreign matter. The origin of the spore-groups is quite remarkable, and I have had some difficulty in making it out, owing to the scantiness of the material which it was essential to injure as little as possible. Their origin seems unassociated with any sexual process, and careful examination of a section shows that the spores, which are practically sessile, origi- nate by budding in all directions from an enlarged hyphal termination. In the fully mature condition which characterizes the type, this termination is very thick-walled and irregular in outline. At points where a spore-group has been cut nearly through the middle, one may see sections of these thick-walled terminations with one or more definitely related spores in situ, as indicated in Figure 34:. Each termination appears to produce as many spores as can be crowded around it, and when the group is viewed from without, it is quite impossible to see any indication of their mode of origin. Although a multiple origin of zygospores from a single conjugation is not neces- sarily excluded as a possibility in this instance, and might find a cer- tain analogy among the Entomophthorales where two distinct zygo- spores may be produced in this manner, it may be assumed that the process in this instance is purely asexual and that it is merely a more specialized manifestation of that which occurs in E. fasciculata, in which, owing to the loose texture of the general mass, the spores, although arising in crowded groups, are produced in a more nearly normal fashion. This conclusion is further supported by the structure of the individual spores which lack a continuous endospore. A majority of the spores are attacked by a sterile parasite similar to that mentioned in the preceding species and shown in the spore at the right in Figure 34. 312 THAXTER. Endogone macrocaepa Tul. Tulasne (1851), p. 182, PL XX, fig. 1. Bucholtz (1912), p. 184, figs. 62-74. Nee Harkness (1889), p. 279. Glomits macrocarpus Tul. (1845), p. 63. Endogone australis Berk. (1860), p. 270. Bucholtz (1912) gives an extended summary of the occurrence and spore-variation in this species, which indicates that it is perhaps the most frequently observed and variable member of the genus. The only records of its occurrence in America are that of Lloyd (1908), who reports it somewhat doubtfully from the Bahamas; and that of Harkness (1899) Avho speaks of finding it under Libocedrus at Towles, in the Sierra Nevada Mountains, California. Mr. Lloyd informs me that the Bahama specimen, which was doubtful, and may have been E. fulra, has been lost; so that this record must remain very dubious. The California form, which I have examined, proves, as above stated, to be E. lactifiua and is identical with what I have called by this name from the East. The spores are clearly zygospores, and the hyphal envelope is well developed, although the "flammenkrone" are not so strikingly differentiated as in some of the Hesse specimens, in the Farlow Herbarium. In New England I first encountered what I have regarded as this species, growing on earth in greenhouse pots at the Botanic Garden in Cambridge, in company with Hymcnogastcr Khischii and Ilydnan- ghnn carneum, a habitat and association which has also been noticed in Europe. Of this material, one gathering made in the winter of 1891-92, has spores seldom exceeding 100 jj. in greatest diameter, while a second gathering made two years later from the same pots, has spore- masses in which the larger spores measure from 170-200 ix in greatest diameter. In neither of these was any definite peridium developed, possibly owing to the fact that both grew on the surface and were subjected to constant watering. In addition to these two gatherings, seven others have been made at Kittery Point, Maine. In these instances the fungus was found in moist coniferous and deciduous woods, usually just below the leaf cover rarely on the surface; the spore-masses usually solitary, or but two or three together. This material also shows a considerable range of variation in the size of the spores; although a majority correspond in this respect to the first gathering above mentioned. The larger spores are in general 80-100 /x in greatest diameter. This average maximum REVISION OF ENDOGONEAE. 313 is considerably below that given by Bucholtz in his summary of the spore measurements of twenty-seven European gatherings; which includes no case in which the maximum is below 100 fj.. When one considers, however, that he gives a variation of the maximum diameter in this summary between 112 ^t and 230 //, the smaller maximum of the American material does not appear significant. The structure and character of the gleba is also subject to variation which bears no evident relation to the size of the spores. The hyphal matrix is thus quite loose in some individuals, and the spore origins correspondingly conspicuous ; while in others it is as densely compacted as in E. laciiflua, so that clearly recognizable spore-origins, though readily made out, have to be sought for. Although Baccarini (1903) has made this difference a basis for the separation of his E. Pampalo- mana (vide infra), it hardly seems a sufficient specific distinction. Through the kindness of Dr. Dodge, I have had an opportunity to examine three gatherings made by Mr. H. E. Parks in California: No. 348 at Saratoga, No. 312 at Aldercroft Creek, and the third at Guadalupe. All of these are unusually well developed. The largest measures 15 mm. dry: the peridium is unusually thick, yellowish white, with adherent humus material. The gleba is firm and dull yello\A'ish in the dry material, although dark brown in the alcoholic specimens. The nearly spherical spores often reach the maximum of 230 fjL mentioned by Bucholtz, and the wall, which may reach a thick- ness of 18 fjL, is traversed by radial canals (?) which, although they are much less strikingly developed in a few other specimens examined in which the walls are unusually thick, are here very numerous and conspicuous, and appear to be associated directly with flattened masses of oily material which adhere to the inner surface, and from the middle point of which they seem to spring. In the absence of intermediate conditions, this California form would be specifically separated from the Eastern ones without question. It seems prefer- able, however, as in the case of E. lactiflna, of which they may prove to be the chlamydosporic condition, to include them under one name until we know more about them. It must be acknowledged, never- theless, that the variations above enumerated may prove too great to justify this procedure, and it is possible that, as in the case of E. lactiflna, in the light of further information, more than one species may emerge from this rather too comprehensive assortment. I am indebted to Miss Wakefield of the Kew Herbarium for an opportunity to examine a portion of the type of E. australis Berkeley, from Tasmania. The spores are like those of E. macrocarpa, the 314 THAXTER. maximum diameter observed being 170 /x. In all its characteristics it comes well within the variations of the present species, and there seems to be no reason for maintaining it as a distinct form. Endogone pampaloniana Baccarini (1903), p. 90, has been examined, through the courtesy of Professor Mattirolo, who has kindly communi- cated a slide of microtome sections from the type of this species. Like most sections of this nature, they are of little use for the pur- poses of specific determination, and it is difficult to decide from them what the distinctive characters, if such exist, really are. Baccarini based the species on the fact that the hyphae between the spores are more copiously developed and compactly woven than in the usual types of E. macrocarpa, in which he conceives the spores, "ampolla," to be simply gregarious, while in E. yampaloniana they form a "cumulo," which he regards as a transitional condition between the loose heap formed in E. macrocarpa, and the more definite sporocarp of E. lactiflua. The different origin of the spores in E. lactiflua would, however, destroy any significance in such a series. The spores corre- spond in size to those of E. macrocarpa, 120-140 n, but have much thinner walls, owing perhaps to the immature condition of the speci- men. As has been mentioned above, similar conditions have been found in New England, although the compact " gleba" is characterized by the usual thick-walled spores, and the same is true of Californian material. Until we have much more information concerning the variations of E. macrocarpa it seems desirable to regard E. pampalon- iana as at best no more than a variety of this species. Endogone tenebrosa nov. sp. (Fig. 46.) Spore-mass spongy, easily disintegrating, blackish. Hyphae loose and friable, 8-40 /x in diameter. Chlamydospores spherical or sub- spherical, 200-270 IX, the largest 260 X 275 ix, brownish yellow, be- coming quite opaque at maturity, the reddish brown wall becoming 15-20 II thick and finally invisible; surrounded by a thin hyaline exospore. In Sphagnum. Little Metis, P. Q. E. C. Jeffrey. The material of this species is so broken up in the fluid in which it is preserved that it is difficult to determine what was the original form of the irregular spongy masses. The huge spores are readily visible with the naked eye, and become absolutely opaque from the darkening REVISION OF ENDOGONEAE. 315 of the contents, and finally of the thick endospore, which, at maturity, is invisible even with bright illumination, and is surrounded by a very thin hyaline exospore. Though sometimes slightly irregular, or slightly longer than broad, they are as a rule rather uniformly and evenly spherical. In structure and development they correspond to those of E. viacrocarpa: but are even more closely comparable with those of the species referred to below, which was found in the stomach of a shrew. Endogone microcarpa Tul. (Figs. 35-37.) Tulasne (1851), p. 182, Plate XX, fig. 2. Bucholtz (1912), p. 192, figs. 75-76. nee Rabh. Fungi Europaei No. 2516. Glomus microcarpus Tulasne (1845), p. 63. This species has been recorded from America only on the authority of Harkness (1899), who collected what he regarded as this form in the forest at ISIill Valley, California, No. 237. The description which he gives does not make at all clear what he had before him ; but the corre- sponding number from the Harkness Collection, which has been kindly sent me for examination b}' the Stanford University Herbarium, proves to correspond to some of the forms of E. lactifiua, the spores being clearly zygospores. A form, however, identical in all respects with the figures and de- scription of Tulasne, has been kindly communicated to me by Dr. C. W. Dodge; who collected it in June, at Aldercroft Creek, Los Gatos, California. The spore-masses are well formed, though rather small, firm and similar to those of E. macrocarpa in form and color. The spores are nearly spherical, 40-48 /x, and very thick-walled. Although there have been various records of this species in Europe, it does not appear, from published accounts, that it has been recog- nized with certainty since the original records of Tulasne, by whom it was found in Italy and France; and it seems to have been confused with smaller types of E. macrocarpa. Some of the latter from America serve in a measure to bridge the gap between the two species, but E. microcarpa, with a rather constant maximum spore diameter of 48 fx, seems clearly distinguished. The accounts of Tulasne and of Buc- holtz, who reexamined the original types, should be consulted for further information in regard to this species. 316 THAXTER. Endogone radiata nov. sp. (Figs. 47-51.) Fruiting body variously lobed, whitish, becoming yellowish brown in alcohol, about 10 X 5 mm., the dried specimen about 5 mm. Gleba tough, dense, nearly homogeneous, the closely coherent rather slender elements hardly distinguishable, yellowish with a fibrous appearance; the peridial layer rather thin, darker brow^nish, the superficial hyphae usually producing terminal and intercalary vesicu- lar enlargements with distinguishing septa. Spores scattered, some- times rather distant, sometimes wdth a slight tendency to grouping, rarely spherical, usually with the longitudinal axis considerably greater than the transverse, oblong, elliptical or subpiriform, often irregular from pressure, the long axis more or less coincident with the radius of the fruiting body, 68 X 38-85 X 50 ju, borne terminally on often clearly recognizable simple hyphae, somewhat stouter than those which compose the substance of the gleba. The spore-wall shows no visible distinction between exospore and endospore and is from 4-5 /x thick: the contents rather finely granular, pale brownish yellow. Under the leaf cover in spruce woods; Gerrish Island, Kittery Point, Maine; Intervale, N. H. ; August, 1896 and 1901 : in Sphagnum, Little Metis, P. Q. E. C. Jeffrey. This species was first taken for E. microcarxM : but is certainly distinct. Its spores are rarely spherical although they appear to be so when cut transversely; the wall is comparatively thin, and is not \'isibly double. The radiate arrangement of the spores, which are firmly embedded in a dense fibrous matrix, seems to be characteristic; but is lost as soon as the section deviates from the radial direction. In the specimens from Kittery and Little Metis, the surface of the peridium shows numerous short projecting filaments with swollen terminations, and intercalary vesicular cells of no great size. At Kittery Point this species was found in company with E. incrassata which was supposed, at the time, to be the same. It is thus not now possible to say whether it had the same alliaceous odor. None was noticed in the Intervale rnaterial. Among the rather numerous individuals collected by Professor Jeft'rey, there are no individuals of E. incrassata, as far as has been ascertained. x\ny connection be- tween the two is thus problematical. REVISION OF ENDOGONEAE. 317 Endogone arenacea nov. sp. (Figs. 38-40.) Spores associated in an indefinite mass through which the material of the substratum (sand) is uniformly and copiously distributed, the whole bound together in an irregular crust-like aggregation, by a loose white mycelium of occasionally septate hyphae. Spores, chlamydospores, rather uniformly spherical, thick-walled, brownish yellow, about 70 fx in diameter (65-75 fj.): the walls 5.5-6.5 /x; with koH, S M. Near margin of brook, Maraval Valley, Port of Spain, Trinidad, B. \Y. I., in sand under trash. This species was found at no great distance from the gathering of E. fulva, hereafter mentioned, from the same locality. The spore- mass has the appearance of a bit of caked sand, about 16 X 15 mm. and about 4 mm. thick when dry. The rather scanty mycelium is visible with a lens over the surface, but it would be unlikely to attract attention, and was preserved and examined almost by accident. The mass is less characteristic and more amorphous than that of any other species, unless it be E. vudtiplcx. The spores, although they show occasional variations in outline and slight differences in size, are exceptionally uniform in these respects as compared with other chlamydosporic types, and are usually quite spherical. The very thick endospore is not continuous, and no septum is present : the thin, often hardly distinguishable, exospore is usually externally roughened by adherent more or less granular disorganized material. The hyphae are much bent and tangled between the spores and sand grains, and the spores often arise from a very short branch. Their non-sexual origin is, however, unquestionable. When treated with potash a rather characteristic smoky stain appears about their insertion. Figure 40. The fatty contents is apt to develop acicular fat crystals, Figures 38, 40. The hyphae show the usual irregularities seen in other species of the genus, and are very rarely septate. Endogone canadensis nov. sp. (Figs. 52-55.) Sporocarp subspherical or irreguhirly lobed; soft, but rather firmly coherent, with a rather well defined whitish (?) peridial layer: gleba dark brown. Spores distinguished by a septum, ovoid to ellipsoid, or 318 THAXTER. somewhat asymmetrical, 70-80 X 54-58 /x very rarely 100 X 65 ^t; the wall hyaline or pale yellowish, 4 ^t thick. Hyphae 8-14 ^i, of the usual type, with occasional clearly defined septa; the sporophores characteristically slender, 5-6 ix. In Sphagnum, Little Metis, P. Q. E. C. Jeffrey. The spore-mass in the material examined, which is all alcoholic, is similar to that of E. radiata. The gleba, however, does not consist of a firm dense matrix in which the spores are firmly held, but is formed of a loose mesh of friable mycelium, of the usual type, in which the spores are free, and are associated with numerous vesicular mostly spherical abortive spores of variable size which eventually shrivel and turn brownish. The species is most nearly related to E. fulva, but is distinguished by its decidedly smaller and more regularly ovoid spores, which are borne on characteristically slender sporophores, and separated by a septum. The nearly hyaline wall of the spore is relatively distinctly thicker; the exospore thin, but rather clearly defined. The fatty coarsely granular contents is at first hyaline, becoming brownish. Endogone borealis nov. sp. (Figs. 44-45.) Spore-mass irregular, coherent, spongy, dark, almost chocolate brown, about seven to eight mm. in greatest diameter. Gleba of loosely woven hyphae, 10-25 /x in diameter, among which much foreign matter and many abortive spores are incorporated. Spores reddish brown, broadly and rather symmetrically elliptical, about 125 X 100 M, the larger 145 X 110 ijl: the thick red-brown walls about 8/x: borne on rather slender hyphae and frequently subtended by a septum. In Sphagnum, Little Metis, P. Q. E. C. Jeffrey. This species seems clearly distinguished by the form and color of its thick-walled spores, the contents of which, in the alcoholic material examined, forms a rather finely granular more or less fibrous proto- plasmic network. It does not seem nearly related to other known species unless it be E. canadensis, from which it is distinguished by the peculiar color and broadly and symmetrically elliptical outline of its large thick-walled spores. The endospore is not continuous when ex- amined under brilliant illumination although the isthmus is a very narrow one and a small septum appears to be present. REVISION OF ENDOGONEAE. 319 EndogoNe pulvinata Henn. (Figs. 41-43.) Hennings (1897), p. 212: nee Lloyd (1918), p. 800, fig. 1240. Dr. Lindau has very kindly allowed me to see a fragment of the type of this species, collected by Gollmer, and found growing on the ground at Caracas, Venezuela. The specimen, which is not in the best condition, resembles E. fulva in general appearance and color. The spores, however, although they have thin walls like E. fulva and are similarly separated from the hypha by a septum, are distinctly different in general appearance, being more nearly spherical, often asymmetrical, and seldom showing the considerable difference l)etween the two diameters that is so characteristic in the last mentioned species. The larger spores are 85 X 85 /i or 75 X 85 ix, according as the axes tend to vary slightly : the average being about 75 X 75 m or 75 X 70 fjL, with considerable variation below these dimensions, and no little variation in outline. The walls are 2-4 jjl thick, as in E. fulva, and the hyphae which, in the specimen seen, are for the most part disorganized, appear to be entirely similar and loosely woven. Endogone fulva (Berk.) Pat. (Figs. 56-59.) Paurocotylis fulva Berkeley (1873), p. 137. Endogone Moelleri Hennings (1897), p. 211. Endogone lignicola Patouillard (1902), p. 183. Bucholtz (1912), p. 199, figs. 97-99. Endogone fulva Patouillard (1903), p. 341; Bucholtz (1912), p. 200, figs. 97-99. Endogone pulvinata Lloyd (1918), p. 800, fig. 1240, nee E. pulvinata Henn., (1897), p. 212. Patouillard first called attention to the fact that Paurocotylis fulva belonged to the genus Endogone and that it was unrelated to P. pila Berk, which is the type of the genus. From the data and figures given by Bucholtz, who has examined the original material in both instances, the identity which he suggests between E. fulva and E. lignicola seems almost certain. The fact that they occur in widely separated regions, the one in Ceylon, the other in the West Indies, is shown to be of little significance; since other species, like E. malleola may have, as will be seen, an equally wide distribution. 320 THAXTER. I have collected this species in abundance in the Maraval Valley near Port of Spain, Trinidad, growing subgregariously along the Maraval brook in moist bamboo trash, fruiting within this material and running out to produce its fructifications on the surrounding sand and pebbles. A single specimen was also found under the leaf cover in the forest about the Grand Etang, Grenada; and I obtained several typical specimens growing exposed on rotten logs in Boggs' Hammock, a short distance south of Cocoanut Grove, Florida. Dr. Lindau has been so kind as to send me a fragment of the type of E. Moelleri, described by Hennings from Brazil. This material is, as above indicated, identical with the Trinidad form, which has been submitted to M. Patouillard and is pronounced by him in all respects the same as his E. lignicola. The spores of the Brazilian form have the darker color which seems to be more characteristic of individuals which have developed in humus, without exposure to the light and air, and are, as in the Grenada gathering, sometimes almost opaque when first mounted. Mr. Lloyd has also been so kind as to send me a portion of the Jamaica material figured by him (1. c.) as E. pidvinata Hennings, as well as a second specimen collected by Mr. Brace in the Bahamas. These gatherings also correspond in all respects to the Trinidad form, and must be regarded as typical E. fulva. I have further received from Professor Mattirolo for examination, a specimen collected by Rick in Brazil, which also has all the essential characters of the present spe- cies, although the spores are not turgescent: and from Professor Spegazzini a gathering from La Boca, Buenos Aires, doubtfully de- termined as A. argentina, which seems quite typical of this species, although not in very good condition. The spore-masses of E. fulva vary from 1^ cm. to a few mm. in diameter when dry, and are usually umbilicate below, subspherical to flattened and irregularly lobed; and even in the same gathering there may be great variation in color. The peridium, which is usually well developed, although in some specimens it may be absent to a greater or less extent, exposing the naked spore-mass, is at first pure white and floccose in young fresh individuals, turning brownish with age, or when handled, the color deepening from ochraceous tawny to chestnut brown. The hyphae are of the usual type, rather stout, 8-12 (x in diameter, more or less, often nodulose or irregular, showing occasional septa, which are more frequent than in most other species, and are in some cases quite loosely interwoven. REVISION OF ENDOGONEAE. 321 The spores vary considerably in color, even in the same indi\'i(hial; and although sometimes nearly opaque, "atro olivaceis vel atris," ma}', when produced free to the light and air, have a decidedly pale, yellowish color. Their outline is characteristically oblong, elliptical to oval or even subpiriform, rarely nearly circular in outline, except when viewed end on. They may be more than twice as long as broad, e.g. 125 X 55 /i, and ordinarily show a decided difference between the long and short diameter; the average variation being from 50- 125 X 45-70 fjL. The wall, although thin as compared with some forms of E. macrocarpa, for example, is thick, 2-4 n, in contrast to the walls of the sporangial types. Bucholtz makes a separate category, a fourth subdivision of the genus, to include this somewhat thinner walled type of spore, and speaks of them as possible sporangia. Hav- ing examined a large series of specimens in all stages of development, and from widely separated localities, it seems evident that they are certainly nothing more than chlamydospores, having somewhat thinner walls than those of the more familiar species, and being distinguished by a septum. The attachment of the spore is often sublateral, as is indicated in figure 97 of Bucholtz, and the sporogenous hypha is often, though by no means invariably, somewhat narrower just below the point of attachment. , The contents of the spores may be rather dense and uniformly granular, or is often somewhat stringy in appearance apparently from the presence of fatty crystalline structures. The species is most nearly related to E. puhinata and the other forms in which the spore is distinguished by a basal septum. Endogone reniformis Bres. (Figs. 60-71.) Bresadola (1896), p. 297. Endogone ? argentina Spegazzini (1899), p. 300. Through the kindness of Professors Lindau and Spegazzini I have been able to examine the type material of E. reniformis Bres. collected by Moller in Brazil and of E. argentina collected at Santa Catalina, Llavallol, Argentina. The Abbe Bresadola has also sent me a third specimen collected by Rick in Brazil, and I myself found apparently the same form in the antarctic forest at Punta Arenas, Magellanes, Chile. 322 THAXTER. A comparison of these four gatherings indicates tliat, although the spores of the Magelhm specimen are distinctly larger, the other three are not separable, and correspond in all essentials. Bresadola, in his description, spealvs of monosporic asci in which the spore is clearly distinguished, but was probably misled by the appearance of young sporangia in which the contents was still continuous, not having yet divided into spores. The sporangiocarps of this species which occur on or just under the leaf cover, are subspherical to reniform, umbilicate, yellowish when dry, nearly white when fresh, 4-10 mm. in diameter, sometimes 20 mm. according to Spegazzini, and arise from a ropy mycelium which may form a more or less distinct stalk as in E. mallcola. In the specimens examined there is no peridial layer distinguishable, the surface being composed of sporangia and slightly projecting scanty hyphal elements. The fertile liyphae are sparingly septate and branched, bearing the sporangia terminally and diverging from a cushion-like basal region associated with the umbilicus. The sj)orangia are more commonly spherical, but, as in E. inalhola may show variations in length and breadth and may be asymmetrical in outline (Figs. 61-62). At maturity the sporangium wall collapses about the spores and follows their irregular contour. The average diameter is about 35-40 fx, but may reach 60 ju or over. The spores, which are evidently formed by cleavage of the whole contents in these sporangia, vary in number from four to a dozen or even more, although Spegazzini mentions eight, only, and are rather variable in size and irregular in shape from mutual pressure. In the Brazilian and Argentine material. Figures 04-71, they are 12-30 X 12-25 fi the average about 18 X 20 n, but in the Magellan material. Figures 60-63, they are for the most part distinctly larger, 20-38 X 14-34 fx. The number present in a single sporangium varies from four to a dozen or more; although, as stated by Spegazzini, there are often not more than eight. This number is, however, by no means constant or even characteristic. On the rupture of the spor- angium wall they are readily set free, although when fully mature, Figures 67-68, they appear to be held by the collapsed sporangial wall and rather firmly coherent. They are quite hyaline and con- tain, as a rule, one or more large oil globules or coarse dense granules. A second Argentine collection from La Plata sent me by Spegazzini doubtfully determined as this species, proves to heE.fulva, as already mentioned. REVISION OF ENDOGONEAE. 323 Endogone malleola Harkn. (Figs. 72-78.) Harkness (1899), p. 280, Plate XLIV, figs. 22 a & b. Endogone microcarpa Fischer pro parte (1897), p. 121, figs. 4-5. Rahenhorst Fungi Europei, No. 2516, nee Tulasne (1851). Endogone pisiformis Bucholtz (1912), p. 196, figs. 88-96; nee Link (1809). E. Torrendii Bresadola. In Torrend (191.3), p. 101: (1920), p. 55. Torrend (1913), Fungi Selecti Exsiccati, No. 1.59. This species seems to have been responsible for much of the con- fusion with which the genus has been afflicted, since, although it is fundamentally unlike the majority of the other types which have been included in Endogone, it bears certain resemblances to them which have led to a misconstruction of appearances that are frequently found in the spores of the other two sections of the genus. This mis- conception has led to the opinion that the chlamydospores, for ex- ample, were to be regarded as sporangia, or at least that they might become directly transformed into sporangia. This conclusion, how- ever, seems to have no better basis than the fact that, in many cases, the contents of these spores is so modified, that they become filled with large granules or fatty bodies, often so uniform in size and form that their spore-like character has been assumed. Thus Bucholtz in his Beitrage, influenced probably by the use of the terms sporangium and sporangiolum in Link's description, has assumed that the present form may be regarded as the true E. -pisiformis, and is thus the type of the genus. The reasons for belie\'ing that this reference can hardly be correct, have already been mentioned. In E. malleola, however, the large spherical or somewhat irregular bodies which form the fructifying mass are filled with numerous relatively large, separable, walled spores; quite different in appearance from any differentiation such as has been above referred to. The references to this species which occur in the literature, are for the most part based on the material collected by Vittadini in the vicin- ity of Naples and distributed in the Fungi Europaei under E. micro- carpa. Fischer (1896) assuming that the determination was correct, and that the material showed a condition of this species in which the chlam\dospores had become transformed into sporangia, regarded it as a demonstration of the sporangial or hemiascoid nature of the spores of Endogone in general. 324 THAXTER. The significance of this condition has been variously discussed, and the terms ascus and sporangium variously applied to it. Its re- semblance to the sporangium of the ]Mortierelleae was first pointed out by Baccarini (1903), who believed that it should be excluded from the Endogoneae for this reason. The researches of Bucholtz who demonstrated the sexual origin of the spores in certain species, and the necessity of their inclusion among the IMucorales, gave further support to this suggestion of Baccarini, and, assuming that the three sections herewith distinguished actually represent conditions of a single generic type, the view that the members of the family are close relatives, at least, of the Mortierelleae, is, as has been already pointed out, strongly supported by the fact that in this family alone among the Mucorales, does one find zygospores ha^^ng specialized envelopes, associated with highly developed acrogenous chlamydospores; and sporangia sepa- rated from the sporangiophore by a simple septum. It should be remembered, however, that although the two may be provisionally thus associated, the apparent parallelism is not necessarily more than a coincidence. The second record of this species is that of Harkness (1899) who first described it under the name E. malleola from material, collected on Mt. Tamalpais in California, which I have had the privilege of ex- amining, and which differs in no essential from the Naples material, although the maximum diameters of the latter are often greater (Figs. 72-74). The form was not again reported till specimens collected in Portugal were described as E. Torrendii Bresadola, Figures 75-76, in an enumer- ation by Torrend (1913) of the second century of his Fungi Selecti Exsiccati, published in Broteria. Quite recently this description has been republished by Bresadola (1920) among his Selecta Mycologica, where, however, the fact of its distribution by Torrend is not men- tioned. Its range has been further extended by its discovery in New Zea- land where material, having dimensions somewhat greater than those of the Naples gathering, has been collected by jNIr. James Mitchell, and very kindly communicated to me by Mr. Lloyd (Figs. 77-78). If one compares these different gatherings, although there is a general agreement in the form, structure and color of the fruiting masses, which are very similar to those of E. arcjcntina, the average size of the sporangia and the number of spores which they contain is subject to considerable variation. Treatment with potash, slight pressure of the coverglass, and degrees of maturity, have to be con- sidered in such a comparison; but quite apart from these, there is a REVISION OF ENDOGONEAE. 325 marked difference observable even between ^^di^•iduals of the same gathering. Thus of two individuals from the Torrend distribution, one shows sporangia with an average diameter of 55-GO /x, while those of the other average from 70-75 jx or slightly over, the latter dimen- sions corresponding to the Californian and Naples gatherings. Al- though Bucholtz reports a maximum diameter of 116 // for the latter, I have not seen any above 100 // in the specimen examined. The New Zealand form, on the other hand, is distinctly larger, the maximum diameter being 120 /x, diameters of 100 ju being common and the average being 80-85 n. The form of the sporangia is normally subspherical, but may be irregular, longer than broad, or even broader than long, or subangular from mutual pressure. The wall usually appears thin, and tends to follow the contour of the contained spores; but, especially when treated with potash, may form a clear gelatinous envelope around the spores, 4-5 jj. thick. The spores are somewhat variable in size, sub- angular from pressure, but often become spherical when free, and possess a distinct thin wall. None have been seen, even in the Tor- rend material, which closely approach the measurements given by Bresadola, 15-28 X 15-17 ju. Measured in the sporangium they rarely seem to exceed 14-15 ix, and usually average from 8-12 fi: although when set free and treated with potash they may reach 20 n occasionally. They form a rather viscous^ mass, and when the sporangium is violently broken, are apt to escape in more or less coherent groups. The filaments, on which the sporangia are borne terminally, are branched and usually rather copiously septate, even submoniliform; the contents above the upper septum, which is often a short distance below the sporangium, being often divided into several superposed spores. From its general characters this form could probably be cultivated with ease by anyone who was fortunate enough to find it in a fresh condition, and a thorough examination of its development in pure cultures is very much to be desired. Doubtful or Excluded Species of Endogone. Reference has been made above to the occurrence of spores of En- dogone in the digestive tract of animals, and in this connection it may be mentioned that in one of these instances spores and mycelium were found in the stomach of a shrew, sent me by Mr. Judd from the vicinity of Washington, D.C. In this material, scanty but typical Endogone filaments bear a few very large spores, some of them 240 /x 326 THAXTER. in diameter, similar to those of E. macrocarpa, when young, but be- coming quite opaque as they mature, owing to a blackening of the exospore. This cannot apparently be referred to any of the described species, although it is very similar to E. tenehrosa. The opacity of the spore, however, seems due rather to the formation of a black encrusta- tion than to a gradual darkening of the contents such as takes place in E. tenehrosa. A second type found in the digestive tract of a m\Tiopod collected in Eastern Tennessee, appears also to belong to an undescribed En- dogone. The hyphae and spores are typical of this genus, the latter browTiish yellow, mostly longer than broad, the greater diameter about 38-45 At, the walls not greatly thickened, peculiar from its slightly one- sided insertion on the sporiferous hyphae. Its size is very near that of E. microcarpa, but it differs in its much thinner wall, asymmetrical insertion and more elongate outline. On the other hand it differs from E. fulva in its smaller spores with relatively thicker walls. A third form, which approaches more nearly to some of the varia- tions of E. viacrocarpa, was observed by Dr. ^Yeston while working with water moulds in the Harvard Laboratory. It produced a rather scanty groT\i;h, consisting of a single subdichotomously branching hypha having all the characteristics of those peculiar to the genus. This grew in water about a fly, attacked by Saprolegniae, and pro- duced abundant spores rather thin-walled, subspherical, pale brownish yellow, the larger 85-100 ix in diameter. It is quite probable that this represents a form of E. macrocarpa, modified by its growth under unnatural conditions. E. Tozziana Sacc. & Cav. has been referred to Leucogaster, a dispo- sition which is confirmed by an examination of a portion of the type. SPHAEROCREAS Sacc. & Ell. Type species Sphaerocreas pubescens Sacc. & Ellis. (Figs. 79-82.) Saccardo & Ellis (1882), p. 582. Stigmatella pvhescens Saccardo (1886), p. 680. Sderocystis pvbescens von Hohnel (1910), p. 399. This species was based on rather scanty material collected on leaves and sticks at Newfield, New Jersey, by Ellis; a portion of which has REVISION OF ENDOGONEAE. 327 been examined in the Farlow Herbarium. For some inexplicable reason it was later associated in the fourth volume of the Sylloge, in the genus Stigmatella, with a second form, Stigmatella aurantiaca B. & C, a wholly different organism belonging to the Myxobacteriaceae, as I have formerly pointed out (Thaxter (1892), p. 402), where the close relationship of S. pubcsccns to Endogone is also referred to. Von Hohnel (1909), p. 127, includes in this genus his own